Raphanus raphanistrum subsp.
sativus
Scientific Name
Raphanus raphanistrum subsp. sativus (L.)
Domin
Synonyms
Raphanistrum gayanum Fisch. & C.A.Mey.,
Raphanus acanthiformis Morel ex L.Sisley,
Raphanus acanthiformis J.M. Morel ex Sasaki,
Raphanus acanthiformis var. raphanistroides
(Makino) Hara, Raphanus candidus Vorosch.,
Raphanus chinensis Mill., Raphanus gayanus
(Fisch. & C.A.Mey.) G.Don, Raphanus macropo-
dus H.Lév., Raphanus niger Mill., Raphanus
oleifer Steud., Raphanus orbicularis Mill.,
Raphanus radicula Pers., Raphanus raphanistroi-
des (Makino) Nakai, Raphanus raphanistrum var.
sativus (L.) Schmalh., Raphanus raphanistrum
subsp. sativus Schmalh., Raphanus raphanistrum
var. sativus (L.) Domin, Raphanus raphanistrum
var. sativus (L.) Beck, Raphanus rotundus Mill.,
Raphanus sativus L., Raphanus sativus subsp.
esculentus Metzg., Raphanus sativus var. longi-
pinnatus L.H.Bailey, Raphanus sativus var. mac-
ropodus (H.Lév.) Makino, Raphanus sativus var.
niger (Mill.) J.Kern., Raphanus sativus var. radic-
ula Pers., Raphanus sativus var. raphanistroides
(Makino) Makino, Raphanus sativus f. raphanis-
troides Makino, Raphanus sativus subsp. sinensis
Sazonova & Stank., Raphanus sinensis Thunb. ex
Pritz., Raphanus taquetii H.Lév.
T.K. Lim, Edible Medicinal and Non Medicinal Plants: Volume 9, Modified Stems, Roots, Bulbs,
DOI 10.1007/978-94-017-9511-1_31, © Springer Science+Business Media Dordrecht 2015
Family
Brassicaceae
Common/English Names
Chinese Radish, Common Radish, Daikon,
Daikon Radish, Fodder Radish, Garden Radish,
Japanese Radish, Leafy Daikon, Oriental Radish,
Oriental Winter Radish, Radish, Wild Radish
Vernacular Names
Arabic: Fejil, Fijil, Fujl
Azerbaijan: Qırmızı turp, Turp
Brazil: Rabanete
Burmese: Monla
Chinese: Hung Loh Paak Tsai, Hong Luo Bo Zi,
Lai Fu, Lai-Fu-Tzu Ts-Ao, Loh Bak, Luo Bo,
Ou Zhou Luo Bo
Croatian: Rotkva, Rotkvica
Czech: Øedkvièka, Ředkev Seta
Danish: Haveræddike, Kinaræddike, Ræddike,
Radis, Radise, Japanræddike
Dutch: Radijs, Ramenas, Tamme Radÿs
Esperanto: Rafano, Rafano Kultiva, Rafano
Manĝebla, Rafaneto
Estonian: Redis
Euskera: Arapa, Erradilla, Erraso, Errasu, Errefau,
Errefaun, Erresan, Erresana, Erresauchoa,
Erresaun, Erresauna, Erresautxo, Lutxarbi
829
830 Brassicaceae

Finnish: Retiisi, Retikka, Ruokaretikka Serbian: Rotkvica, Trotkvica

French: Petit Radis, Radis, Rave, Radis Cultivé, Slovak: Reďkev, Redkev Vrtna, Redkvica, Vrtna
Radis D’été, Radis Rose Redkev
Galician: Labestro, Rábano, Rabâo, Rábâo, Slovenian: Retvica
Ravo, Saramago Spanish: Erradil, Labrestos, Matacandil,
German: Bierrettich, Bierwurz, Furzwurzel, Rabanillo, Rabanito, Rabaniza, Rábano,
Garten-Rettich, Monatsrettich, Radi, Rábano Blanco, Rábano Blanco Redondo,
Radieschen, Rettich, Retwurzel Rábano Castellano, Rábano Colorado
Greek: Rapani Redondo, Rábano Común, Rábano Encarnado,
Hebrew: Tznonit Rábano Granadino, Rábano Largo, Rábano
Hornjoserbsce (Upper Sorbian): Zahrodna Macho, Rábano Pajizo, Rábano Redondo,
Rjetkej Rábano Silvestre, Rábanos, Zanahoria
Hungarian: Retek Sri Lanka: Rabu (Sinhalese)
Icelandic: Ætihreòka, Hreðka, Raefla Swedish: Rädisa
India: Mulō, Mulla (Bengali), Mooli, Muli, Thai: Hua Phak Kat Khao, Hua Pàk Gàat Kaao,
Mūlī, Mulla (Hindi), Mullangi (Malayalam), Hua Chai Táo, Ma Puek (Chiang Mai), Phak
Mūlaka (Marathi), Mūlī (Punjabi), Muulaka Kat Hua (Central), Phak Poek Hua
(Sanskrit), Muulam, Mullangi, Mullanggki (Northern)
(Tamil), Mullangi (Telugu), Mūlī (Urdu) Turkish: Trup, Tūrp
Indonesia: Lobak, Lobak Berem, Lobak Putih, Vietnamese: Củ Cải, Củ Dền, Rađi
Luba (Malay) Lobak, Rades (Javanese), Welsh: Heddig, Radys, Redis, Redyns, Rhodri
Lobak Berem, (Sundanese) Rhuddygl
Italian: Rafano, Ramolaccio, Ravanello,
Ravanello Commune
Japanese: Daikon, Hatsuka Daikon, Radeisshu Origin/Distribution
Khmer: Chhaay Thaw
Kurdish: Tūr The exact origin or radish is unknown because it
Korean: Il Mu, Mu has been in cultivation for thousands of years.
Laotian: Kaad Khaaw Radish is thought to have originated in the east-
Latvian: Redīss ern Mediterranean region and has been domesti-
Lithuanian: Valgomasis Ridikas cated in Europe in the pre-Roman times.
Malaysia: Lobak, Lobak Putih, Lobak Merah, DNA sequence study from different organ-
Luba elles by Yang et al. (2000) suggested Raphanus
Nepalese: Mulo sativus to be a hybrid between B. rapa/B. olera-
Norwegian: Hagereddik, Reddik cea and B. nigra lineages as proposed by Song
Persian: Torobcheh et al. (1990).
Polish: Rzodkiew, Rzodkiewka
Philippines: Rabanos (Cebu Bisaya), Labanos
(Tagalog) Agroecology
Portuguese: Raba, Rabanete, Rabanetes
Escarlate, Rábano, Rabão, Rabâo Radisio, Radish tolerates a wide range of climatic
Rabiça, Rabiças, Rabo regimes. Radish flourishes in full sun and in
Romanian: Ridiche well-drained, light sandy loams with pH of 6.5–
Russian: Red’ka, Red’ka Ogorodnaia, Red’ka 7. In temperate areas, radish is usually planted
Posevnaia out a few weeks prior to spring before the first
Sardinian: Ravanella frost. In warm weather, they are normally planted
Scots: Reefort out in fall.
Raphanus raphanistrum subsp. sativus 831

Plate 1 (a, b) Chinese radish (lobak)

Edible Plant Parts and Uses

Radish roots, seed sprouts, leaves, flowers and

seeds are edible. Radish roots are eaten raw,
baked, roasted or cooked in stews, soups or stir
fries, also lightly steamed with olive oil, salt or
lemon juice. Radish is eaten raw with a dip or
with peanut butter, shredded to salads, pickled
in kimchi and made into radish cakes like
Japanese diakon mocha. A number of pickled
radish products are produced in Korea such as
kaktugi, tongchimi, chonggak-kimchi, seok-
bakji, yolmu-kimchi dan moogi kach doo ki gac-
tuki and mootsanji. Sinki is a sour pickle
prepared from radish taproots and is consumed
traditionally in India, Nepal and parts of Bhutan,
where it is used as a base for soup or eaten as a
pickle.
Young leaves are also eaten raw or cooked as
greens. Radish greens are also blended into
smoothies. The blossoms are edible, and the seed
pods have many uses, eaten raw, cooked, in stir
fries to pickles.
Botany
An annual or biennial coarse herb, 30–120 cm
tall, glabrous, scabrous or hispid with stout, erect,
branched stem. Roots are fleshy; swollen; white,
pink, red or black; linear; fusiform; oblong; or
Plate 2 Daikon plant
globose, 1–100 × 0.5–45 cm, sometimes slender
and not fleshy (Plates 1, 2, 3, 4 and 5). Lower
leaves are lyrate-pinnatifid, petiolate, sparsely
hirsute to glabrous; the lobes are dentate or den-
ticulate (Plates 2 and 6). The uppermost cauline
leaves are subsessile, often undivided and den-
tate. Racemes are elongated, narrowly erect,
oblong, glabrous or sparsely hirsute; petals are
purplish, pink to nearly white (Plate 7), with dark
purple veins, 1.5–2 cm long, broadly obovate
with a long narrowed claw; filaments are slender
and anthers are sagittate at the base, style, stigma
entire. Fruit is indehiscent, silique fusiform, lan-
ceolate and cylindrical to conical, tapering from
above base to a narrow beak. Seeds are 2–4, glo-
bose or ovoid, somewhat ridged, 2.5–4 mm in
diameter.
832 Brassicaceae

Plate 6 Radish leaves

Plate 3 Daikon radish

Plate 7 Radish flowers

Nutritive/Medicinal Properties
Plate 4 European radish with small short cylindrical red
roots Nutrients/Phytochemicals: Root

The proximate nutrient composition per 100 g

Plate 5 European radish with small globose red roots
edible portion of raw radish was reported as
water 95.2 g, energy 16 kcal (66 kJ), protein
0.68 g, total lipid 0.10 g, ash 0.55 g, carbohydrate
3.4 g and total dietary fibre 1.6; total sugars
1.86 g, sucrose 0.10 g, glucose 1.05 g and fruc-
tose 0.71 g; minerals, Ca 25 mg, Fe 0.34 mg, Mg
10 mg, P 20 mg, K 233 mg, Na 39 mg, Zn
0.28 mg, Cu 0.050 mg, Mn 0.069 mg and Se
0.6 μg; vitamins, vitamin C 14.8 mg, thiamine
0.012 mg, riboflavin 0.039 mg, niacin 0.254 mg,
pantothenic acid 0.165 mg, vitamin B6 0.071 mg,
total folate 25 μg, total choline 12.3 mg,
β-carotene 4 μg, vitamin A 7 IU, vitamin K (phyl-
loquinone) 1.3 μg, β-carotene 22 μg, lutein + zea-
xanthin 10 μg and phytosterols 7 mg; total
Raphanus raphanistrum subsp. sativus
saturated fatty acids 0.032 g, 16:0 (palmitic acid)
0.027 g, and 18:0 (stearic acid) 0.004 g; total
monounsaturated fatty acids 0.017 g, 18:1 undif-
ferentiated (oleic acid) 0.017 g; and total polyun-
saturated fatty acids 0.048 g, 18:2 undifferentiated
(linoleic acid) 0.017 g and 18:3 undifferentiated
(linolenic acid) 0.031 g; and amino acids, trypto-
phan 0.009 g, threonine 0.023 g, isoleucine
0.020 g, leucine 0.031 g, lysine 0.033 g, methio-
nine 0.010 g, cystine 0.010 g, phenylalanine
0.036 g, tyrosine 0.009 g, valine 0.035 g, arginine
0.038 g, histidine 0.013 g, alanine 0.26 g, aspartic
acid 0.064 g, glutamic acid 0.157 g, glycine
0.026 g, proline 0.022 g and serine 0.278 g
(USDA ARS 2013). Radish was also found to
contain 0.4–2.11 mg/100 g FW of the flavonol
kaempferol FW (Bilyk and Sapers 1985; Hertog
et al. 1992; Lugasi and Hovari 2000). Radish
leaves and sprouts were found to contain
7.72 mg/100 g and 13.76–35.128 72 mg/100 g
kaempferol, respectively (Sakakibara et al. 2003).
The proximate value per 100 g edible portion
of raw oriental radish was reported as water
94.62 g, energy 18 kcal (76 kJ), protein 0.60 g,
total lipid 0.10 g, ash 0.58 g, carbohydrate 4.1 g,
total dietary fibre 1.6 and total sugars 2.5 g; min-
erals, Ca 27 mg, Fe 0.4 mg, Mg 16 mg, P 23 mg,
K 227 mg, Na 21 mg, Zn 0.15 mg, Cu 0.115 mg,
Mn 0.038 mg and Se 0.7 μg; vitamins, vitamin C
22 mg, thiamine 0.02 mg, riboflavin 0.02 mg,
niacin 0.2 mg, pantothenic acid 0.138 mg, vita-
min B6 0.046 mg, total folate 7.3 μg, betaine
0.1 mg, total choline 12.3 mg and vitamin K
(phylloquinone) 0.3 μg; total saturated fatty acids
0.03 g, 16:0 (palmitic acid) 0.026 g and 18:0
(stearic acid) 0.004 g; total monounsaturated
fatty acids 0.017 g, 18:1 undifferentiated (oleic
acid) 0.017 g; total polyunsaturated fatty acids
0.045 g, 18:2 undifferentiated (linoleic acid)
0.016 g and 18:3 undifferentiated (linolenic acid)
0.029 g; and amino acids, tryptophan 0.003 g,
threonine 0.025 g,, isoleucine 0.026 g, leucine
0.031 g, lysine 0.030 g, methionine 0.006 g, cys-
tine 0.005 g, phenylalanine 0.020 g, tyrosine
0.011 g, valine 0.028 g, arginine 0.035 g, histi-
dine 0.011 g, alanine 0.19 g, aspartic acid 0.041 g,
glutamic acid 0.113 g, glycine 0.019 g, proline
0.015 g and serine 0.018 g (USDA ARS 2013).
833
Among the seven Japanese radish cultivars,
Koshin, Kouto and Shogoin were the three high-
est in terms of the total soluble sugar (glucose,
fructose, sucrose) content (Hara et al. 2011).
Sobutori, which is the most common radish culti-
var in Japan, was the lowest. The total organic
acid (malate, citrate, ascorbate, lactate and pyru-
vate) contents varied among the seven cultivars,
although they were 5–13 times lower than the
total soluble sugar contents. The Koshin cultivar
showed the highest amylase activity, with a level
approximately six times higher than that of the
Sobutori cultivar, which had the lowest (Hara
et al. 2009). Cultivars with higher amylase activi-
ties showed higher starch contents.
Anthocyanins
The anthocyanin concentrations of the mature rad-
ish taproot were significantly greater than in the
sprouts of red, pink and purple varieties. The pri-
mary anthocyanidins present in the red and pink
radish varieties were pelargonidin and delphinidin,
while the primary anthocyanidin in the purple rad-
ish variety was cyanidin (Hanlon and Barnes 2011).
Radish was reported to contain 7.40–128 mg/100 g
pelargonidin anthocyanidins (Harnly et al. 2006;
Wu et al. 2006). Two novel diacylated anthocya-
nins, pelargonidin 3-O-[2-O-(β-glucopyranosyl)-
6- O -( trans - p -coumaroyl)-β- glucopyranoside]
5-O-(6-O-malonyl-β-glucopyranoside) and
pelargonidin 3- O -[2- O -(β-glucopyranosyl)-
6-O-(trans-feruloyl)-β-glucopyranoside] 5-O-
(6-O-malonyl-β-glucopyranoside), were character-
ised from red radish (Guisti et al. 1998). Two other
monoacylated anthocyanins were determined to
be pelargonidin 3-O-[2-O-(β-glucopyranosyl)-6-
O -( trans - p -coumaroyl)-β- D - glucopyranoside]
5- O -(β-glucopyranoside) and pelargonidin
3- O -[2- O -(β-glucopyranosyl)-6- O -( trans -
feruloyl)-β-glucopyranoside] 5-O-(β-glucopy-
ranoside). Twelve acylated anthocyanins were
isolated from the red radish (Otsuki et al. 2002).
Six of these were identified as pelargonidin 3-O-[6-
O-(E)-feruloyl-2-O-β-D-glucopyranosyl]-(1→2)-
β-D-glucopyranoside]-5-O-(β-D-glucopyranoside);
pelargonidin 3-O-[6-O-(E)-caffeoyl-2-O-(6-(E)-
feruloyl-β- D - glucopyranosyl )-(1→2)-β- D -
glucopyranoside]-5-O-(β-D-glucopyranoside);
834
pelargonidin 3-O-[6-O-(E)-p-coumaroyl-2-O-(6-
( E )-caffeoyl-β- D -glucopyranosyl)-(1→2)-β- D -
glucopyranoside]-5-O-(β-D-glucopyranoside);
pelargonidin 3-O-[6-O-(E)-feruloyl-2-O-(6-(E)-
caffeoyl-β- D -glucopyranosyl)-(1→2)-β- D - glu-
copyranoside]-5- O -(β- D -glucopyranoside);
pelargonidin 3-O-[6-O-(E)-p-coumaroyl-2-O-(6-
( E )-feruloyl-β- D -glucopyranosyl)-(1→2)-β- D -
glucopyranoside]-5-O-(β-D-glucopyranoside);
and pelargonidin 3-O-[6-O-(E)-feruloyl-2-O-(2-
( E )-feruloyl-β- D -glucopyranosyl)-(1→2)-β- D -
glucopyranoside]-5-O-(β-D-glucopyranoside). Ten
known acylated anthocyanins were isolated from
red radish (Liu et al. 2008). The acylated anthocya-
nins are all based on pelargonidin 3-sophoroside-
5-glucoside, acylated with caffeoyl, feruloyl or
p-coumaroyl moieties. The anthocyanins were
pelargonidin 3- O -[6- O -( E )-caffeoyl-2- O -β-
D -glucopyranosyl]-(1→2)-β-D-glucopyranoside)-
5-O-(β-D-glucopyranoside); pelargonidin 3-O-
[6- O -( E )-caffeoyl-2- O -(6-( E )-caffeoyl-β- D -
glucopyranosyl]-(1→2)-β-D-glucopypranoside]-
5-O-(β-D-glucopyranoside); pelargonidin 3-O-
[6- O -( E )-caffeoyl-2- O -(6-( E )-feruloyl-β- D -
glucopyranosyl]-(1→2)-β-D-glucopyranoside]-5-
O-(β-D-glucopyranoside); pelargonidin 3-O-
[6-O-(E)-p-coumaroyl-2-O-β-D-glucopyranosyl]-(1→2)-
β-D-glucopyranoside]-5-O-(β-D-glucopyranoside);
pelargonidin 3-O-[6-O-(E)-p-coumaroyl -2-O-(6-
( E )-caffeoyl-β- D -glucopyranosyl)-(1→2)-β- D -
glucopyranoside]-5-O-(β-D-glucopyranoside);
pelargonidin 3-O-[6-O-(E)-feruloyl-2-O-β-D-glu-
copyranosyl ]-(1→2)-β- D -glucopyranoside]-5-
O-(β-D-glucopyranoside); pelargonidin 3-O-[6-
O -( E )-feruloyl-2- O -(6-( E )-caffeoyl-β- D -
glucopyranosyl)-(1→2)-β-D-glucopyranoside]-5-
O-(β-D-glucopyranoside); pelargonidin 3-O-[6-
O -( E )-p-coumaroyl-2- O -(6-( E )-feruloyl-β- D -
glucopyranosyl)-(1→2)-β-D-glucopyranoside]-
5-O-(β-D-glucopyranoside); pelargonidin 3-O-
[6- O -( E )-feruloyl-2- O -(6-(E)-feruloyl-β- D -
glucopyranosyl)-(1→2)-β-D-glucopyranoside]-5-
O-(β-D-glucopyranoside); and pelargonidin
3-O-[6-O-(E)-feruloyl-2-O-(2-(E)-feruloyl-β-D-
glucopyranosyl)-(1→2)-β-D-glucopyranoside]-5-
O-(β-D-glucopyranoside).
Three new acylated pelargonidin 3-sophoro-
side-5-glucosides were isolated from the root
Brassicaceae
peels, petioles and flowers of red radish,
Raphanus sativus ‘Cherry Mate’, in addition to
the five known anthocyanins, namely,
pelargonidin 3-sophoroside-5-glucoside, pelar-
gonidin 3-[2-(glucosyl)-6-(trans-p-coumaroyl)-
glucoside]-5-glucoside, pelargonidin 3-[2-
(glucosyl)-6-(trans-feruloyl)-glucoside]-5-glu-
coside, pelargonidin 3-[2-(glucosyl)-6-(trans-p-
coumaroyl)-glucoside]-5-(6-malonylglucoside)
and pelargonidin 3-[2-(glucosyl)-6-(trans-
feruloyl)-glucoside]-5-(6-malonylglucoside)
(Tatsuzawa et al. 2008). The structures of the
three new acylated anthocyanins were shown to
be pelargonidin 3-O-[2-O-(β-D-glucopyranosyl)-
6- O -( trans -caffeoyl)- β - D - glucopyranoside ]-
5-O -(6-O-malonyl-β-D-glucopyranoside), its
demalonyl derivative, and pelargonidin 3-O-[2-
O-(β-D-glucopyranosyl)-6-O-(cis-p-coumaroyl)-
β -D -glucopyranoside]-5-O -(6-O -malonyl-β - D -
glucopyranoside). p-Coumaroyl anthocyanins
were the main pigments found in the root, peti-
oles and flowers. Although the trans-p-coumar-
oyl form was abundant in all three plant organs,
its cis form was present in very low amount
within the root but in large amount in the flowers
and petioles. Four new acylated cyanidin glyco-
sides were isolated from the purple roots of
Raphanus sativus ‘Benikanmi’, along with five
known anthocyanins (Tatsuzawa et al. 2010).
Two of these pigments were determined to be
cyanidin 3-O-[2-O-(β-glucopyranosyl)-6-O-
(trans-feruloyl)-β-glucopyranoside]-5-O-[6-O-
(malonyl)-β-glucopyranoside] and cyanidin
3-[2-(glucosyl)-6-(cis-p-coumaroyl)-glucoside]-
5-[6-(malonyl)-glucoside]. Two other new
pigments were obtained in small quantities
and were assigned to be malonyl cyanidin
3-sophoroside-5-glucoside and malonyl cyanidin
3-[2-(glucosyl)-6-(trans-caffeoyl)-glucoside]-
5-glucoside.
Six new acylated anthocyanins were isolated
along with the three known congeners from the
fresh roots of red radishes (Tamura et al. 2010).
Their chemical structures were elucidated as
3-O-[6-O-(E)-caffeoyl-2-O-{6-O-(E)-caffeoyl-
β-D-glucopyranosyl}-β-D-glucopyranosyl]-5-O-
(6-O-malonyl-β-D-glucopyranosyl)pelargonidin;
3- O -[6- O -( E )- p -coumaroyl-2- O -{6- O -( E )-
Raphanus raphanistrum subsp. sativus
caffeoyl-β- D -glucopyranosyl}-β- D - glucopy-
ranosyl]-5-O-(6-O-malonyl-β-D-glucopyranosyl)
pelargonidin;3-O-[6-O-(E)-feruloyl-2-O-{6-O-(E)-p-
coumaroyl-β-D-glucopyranosyl}-β-D-glucopy-
ranosyl]-5- O -β- D -lucopyranosylpelargonidin;
3-O-[6-O-(E)-feruloyl-2-O-{6-O-(E)-caffeoyl-β-
D-glucopyranosyl}-β-D-glucopyranosyl]-5-O-(6-
O - malonyl-β- D -glucopyranosyl)pelargonidin;
3- O -[6- O -( E )- p -coumaroyl-2- O -{6- O -( E )- p -
coumaroyl-β-D-glucopyranosyl}-β-D-glucopy-
ranosyl]-5-O-(6-O-malonyl-β-D-glucopyranosyl)
pelargonidin; and 3-O-[6-O-(E)-feruloyl-2-O-
{6- O -( E )- p -coumaroyl - β - D -glucopyranosyl}-
β- D - glucopyranosyl]-5- O -(6- O -malonyl-β- D -
glucopyranosyl)pelargonidin. Sixty anthocya-
nins, 14 acylated cyanidin 3-(glucosylacyl)
acylsophoroside-5-diglucosides, 24 acylated
cyanidin 3-sophoroside-5-diglucosides and 22
acylated cyanidin 3-sophoroside-5- glucosides,
were found in purple Bordeaux radish (Lin
et al. 2011). The presence of 38 acylated cyani-
din 3-sophoroside-5-diglucosides and around
ten acylated cyanidin 3-sophoroside-5-
malonylglucosides was found reported in R. sati-
vus for the first time.
The light irradiation (fluorescence light,
5,000 lx; at 25 °C) indicated that the red radish
extract (in which major anthocyanins were pelar-
gonidin glycosides acylated with a combination
of p-coumaric, ferulic or caffeic acids) was more
stable at lower pH than at higher pH (Matsufuji
et al. 2007). The HPLC analyses revealed that
diacylated anthocyanins in the extract were more
stable to light at pH 3 than monoacylated antho-
cyanins. No significant difference in degradation
rates of acylated anthocyanins at pH 5 was
observed, whereas anthocyanins acylated with
p-coumaric or ferulic acids were more stable at
pH 7 than ones with caffeic acids. The stability to
heat (at 90–95 °C) showed a tendency similar to
that for light. The degradation behaviour of red
radish extract to H2O2 was almost the same to
those of light and heat, depending on the
pH. However, HPLC analyses revealed that the
stability of individual acylated anthocyanins was
independent of the pH. DPPH radical scavenging
activity of all acylated anthocyanins was higher
than those of pelargonidin and pelargonidin
835
3-glucoside. The activity of acylated anthocyanins
mostly depended on the activity of intramolecu-
lar acyl units (caffeic acid > ferulic acid > p-cou-
maric acid). In another study, 17 pigments were
tentatively identified as pelargonidin 3-sophoro-
side-5-glucoside derivatives with multiple acyla-
tion of hydroxycinnamic acids in the roots of
Chinese radish cultivars (Jing et al. 2012). These
Chinese radish cultivars showed high variation in
anthocyanins (63.77–160.74 mg/100 g FW). A
bright colour (CIELab) of radish anthocyanins
was shown at a wide pH range, comparatively
stable at pH < 4.2. Those anthocyanins also
showed a remarkable thermal stability, following
a zero-order kinetics at pH 2.5 with half-lives of
14.5 or 8.7 hours at 90 or 100 °C, respectively.
Additionally, those cultivars varied in glucosino-
late contents (59.69–163.91 mg/100 g FW),
whereas their degradation was sensitive to pH
and followed a first-order kinetics at pH 5.8 with
half-lives of 11.44 or 7.05 hours at 90 or 100 °C,
respectively. However, the stable pH ranges for
anthocyanins and glucosinolates were different:
pH < 4.2 and pH > 3.6, respectively. In a radish
juice model (pH 5.8/2.5), thermal degradation of
anthocyanins or glucosinolates was associated
closely with media pH values.
Red radish anthocyanins (RRAs) in various
fruit juice beverages (apple, grape, peach, pear,
pomegranate and lemon) displayed a much faster
degradation rate during storage at room tempera-
ture (t1/2 value ≤84.0 days) than did in refriger-
ated temperature (Liu et al. 2014). During
heating, RRAs in peach and pomegranate showed
higher stability than others at these temperatures.
There was a positive correlation (R2 > 0.9128)
between ascorbic acid content of juice beverages
(8–36 mg/100 mg) and stability of RRAs at
70–90 °C. It was found that RRAs in apple and
pear juice beverages were more stable than in
other juice beverages.
Glucosinolates/Isothiocyanates/Nitriles
Intact roots of 109 radish cultivars including oil
radishes (subsp. oleifera) and food radishes
(subsp. radicola) from European, European-
American, Japanese and Korean markets were
found to contain primarily 4-methylthio-3-butenyl
836
glucosinolate (GS) with small amounts of
4-methylsulfinylbutyl GS, 4-methylsulfinyl-3-
butenyl GS and 3-indolylmethyl GS (Carlson
et al. 1985). Regarding total GSs, 80 % or more
of the red European-American radishes had
100–199 μmole/100 g, the Korean 100–
299 μmole/100 g and the Japanese
200–399 μmole/100 g. No correlation was found
between root size and 4-methylthio-3-butenyl-,
3-indolylmethyl- or total GSs. Japanese radish
peelings contained significantly greater concen-
trations of these three constituents than did the
peeled root. Early radish (Rex and Ostergruss
Różowa) contained the least amount of thiocya-
nate compounds and winter radish cv. Murzynka
the highest amount (Capecka 1998). The content
of thiocyanates in leaves was three to five times
higher than that in hypocotyl roots in all cultivars
including Japanese radish (Tokinashi and
Minowase Summer Cross F1) and winter radish
(Monachijska Biała and Murzynka). The changes
in the thiocyanate content during root growth
showed a constant rising tendency in the case of
the leaves of all cultivars and the storage organs
of Murzynka. Elivra and Tatiana (2012) reported
radish to contain thioglycosides sinalbin, sini-
grin and glucobrassicin. They found black radish
to have higher content of isothiocyanates
37.6 mg/100 g versus green radish 33.7 mg/100 g
and higher level of indole compounds
35.91 mg/100 versus 24.41 mg/100 g in green
radish.
Trans-4-(methylthio)-3-butenyl glucosinolate
(4MTBGLS) was isolated as a potassium salt
from radish roots (Visentin et al. 1992). The con-
tents of 4-methylthio-3-butenyl isothiocyanate
(MTBITC) in more than half of the commercial
Japanese radish cultivars ranged from 200 to
300 μmol/100 mL juice (Okano et al. 1990).
Among the cultivars examined, ‘Karami’ showed
the highest content of MTBITC, reaching as much
as 1,735 μmol/100 mL, followed by the ‘Shinshu-
jidaikon’ group of 400–700 μmol/100 mL. The
‘Ninengo’ and ‘Shiroagari’ groups also exhibited
relatively high content averaging more than
300 μmol/100 mL. On the contrary, the
‘Miyashige’ group or Chinese cultivars contained
the lowest level of MTBITC ranging from 100 to
Brassicaceae
200 μmol/100 mL. The result of sensory evalua-
tion on grated radish showed that a cultivar with
higher content of MTBITC was proportionately
more pungent, supporting the notion that MTBITC
was the pungent principle in roots of Japanese
radish.
Studies by Ishii and Saijo (1987) found that
isothiocyanate (ITC) in daikon roots decreased
with days after sowing. The roots harvested in
early summer contained a higher level of ITC
than those harvested in autumn. ITC was higher
in radish roots grown in alluvial soils. Mulching
treatment with plastic film increased ITC in roots.
ITC in the roots increased linearly with increas-
ing sulfate levels in pots filled with vermiculite.
Planting density did not affect ITC in roots. The
range and mean of 4-methylthio-3-butenyl gluco-
sinolate (MTBGLS) contents in 20 daikon culti-
vars were 42–345 μmo1/100 g of fresh weight
and 210 μmol/100 g of fresh weight, respectively
(Ishii et al. 1989b). Among the cultivars exam-
ined, the highest in total glucosinolate content
contained 2.3 times higher than the lowest. The
mean content of MTBGLS corresponded to about
80 % of that of total glucosinolate. 4-Methylthio-
3-butenyl glucosinolate, the predominant gluco-
sinolate in daikon root, was quantitatively
determined after myrosinase enzymatic conver-
sion to its isothiocyanate (Ishii et al. 1989a).
The major volatile components in the oil of
the roots of black, white and red radish in respec-
tive sequence were hexadecanoic acid (palmitic
acid) (32.2, 30.3, 49.9 %), methyl linolenate
(21.7, 13.7, 8.5 %), 4-(methylthio)butyl isothio-
cyanate (erucin) (21.5, 25.7, 17.9 %),
5-(methylthio)-4-pentenenitrile (6.9, 5.3, tr%),
dimethyl trisulfide (1.1, 1.3, 3.8 %), 5-(methyl-
thio)pentyl isothiocyanate (berteroin) (0.5, 2.0,
0.6 %), 4-(methylthio)-3-butenyl isothiocyanate
(0.2, 1.7, 0.7 %) and 2-phenylethyl isothiocya-
nate (1.0, 1.5, 0.1 %) (Blažević and Mastelić
2009). Other isothiocyanates and nitriles included
4-methylpentyl isothiocyanate (0.1, 0.1, 0 %),
benzenepropanenitrile (tr, tr, 0 %), 3-(methyl-
thio)propyl isothiocyanate (iberverin) (tr, 0.5,
0.7 %) and benzyl isothiocyanate (tr, 0.5, tr %).
Alkanes identified in the roots were undecane
(0.1, 0.3, tr%), dodecane (0,1.1,1.4 %), tridecane
Raphanus raphanistrum subsp. sativus
(0.4, 0.5, 1.2 %), tetradecane (tr, 0, 0.7 %), hene-
icosane (0, 2.5, 0 %), tricosane (0, 2.3, 0 %) and
tetracosane (0,1.2, 0 %). Other aliphatic alcohols
and carbonyls identified in the roots were (Z)-3-
hexen-1-ol (tr, 0.7, 0 %), (E,E)-2,4-heptadienal
(0.1,0, 0.1 %), 2-phenylacetaldehyde (0.1, tr,
0 %), 4-ethylbenzaldehyde (0, tr, 0 %),
6,10,14-trimethyl-2-pentadecanone (0.1, 0, 0 %),
isobutyl phthalate (0, 0.1, 0.3 %) and dibutyl
phthalate (0.1, 0.7, 1.1 %). Other fatty acids and
esters identified were (Z)-3-hexenyl acetate (tr, tr,
tr%), dodecanoic acid (lauric acid) (tr, 0, o.5 %),
benzyl benzoate (0.1,0,0 %), tetradecanoic acid
(myristic acid) (1.9,0.5, 0 %) and pentadecanoic
acid (0.2,0, 0 %). Other sulfur and/or nitrogen
compounds identified were dimethyl disulfide
(1.8, tr,1.3 %), 2-acetylthiazole (tr, 0, 0 %),
1-(methylthio)-3-pentanone (0.1, 0.3, 0 %) and
dimethyl tetrasulfide (0, 0, 0.5 %). Terpenes iden-
tified were β-caryophyllene (tr, 0, tr) and neophy-
tadiene (0.1, tr, 0 %). Miscellaneous compound
identified included 4-vinyl-2-methoxyphenol (tr,
0, tr%).
Glucoraphasatin an atypical glucosinolate
mainly found in Raphanus sativus roots and
sprouts and its corresponding isothiocyanate,
4-methylsulfanyl-3-butenyl isothiocyanate, had
been found to upregulate enzymes involved in the
detoxification of carcinogens with the potential
to be used as chemopreventive agents (Montaut
et al. 2010). Six glucosinolates were identified
and quantified in the black radish-based dietary
supplements: glucoraphasatin (0.2–0.48 mg/g),
glucosisaustricin (0.37–0.91 mg/g), glucora-
phenin (0.84–1.27 mg/g), glucoputrajivin (0.14–
0.28 mg/g), glucosisymbrin (0.70–0.99 mg/g)
and gluconasturtiin (0.06–0.12 mg/g) (Ediage
et al. 2011). Glucoraphenin was the most abun-
dant glucosinolate in all samples.
Four radish glucosinolates (glucoraphenin,
dehydroerucin, glucobrassicin and glucoerucin)
were identified from root extracts, and dehydroe-
rucin was found to be the major glucosinolate in
red radish roots (Gao et al. 2014). Application of
chitosan with 76 %, 83 % or 89 % deacetylation
in radish extracts attributed to 26 %, 35 % or
43 % adsorption rate for glucosinolates and 28 %,
26 % or 22 % for anthocyanins, respectively. It
837
was found that the concentration of volatile com-
pounds decreased by 70 %, resulting in the loss
of odorous compounds. The changes in chitosan
spectra before/after adsorption and after desorp-
tion at 1,590 and 3,360 cm(−1) and at broad
bands from 2,600 to 2,000 cm(–1) suggested that
the dominant adsorption mechanisms of gluco-
sinolates on chitosan may be electrostatic attrac-
tions, including hydrogen bonds and charge
neutralisation.
The steam-volatile constituents of fresh rad-
ish of Japanese and Kenyan origin identified
included 4-methylthio-3- butenyl isothiocyanate
and its parent glucosinolate, dimethyl disulfide,
methyl methanethiol sulfinate, pentyl isothiocy-
anate, 4-methylpentyl isothiocyanate, hexyl iso-
thiocyanate, 5-(methylthio)-4-pentenenitrile,
3-methylthiopropyl isothiocyanate, benzyl iso-
thiocyanate, 4-methylthiobutyl isothiocyanate,
5-methylthiopentyl isothiocyanate, (E )-2-
hexenal, (Z)-3-hexenol, 1-methylthio-3-penta-
none, 5-hexenyl isothiocyanate, 5-(methylthio)-
pentane nitrile and 2-phenylethyl isothiocyanate
(Kjaer et al. 1978).
The six heirloom Japanese radish varieties pro-
duced 2.0–11.5 times higher levels of 4-methyl-
thio-3-butenyl isothiocyanate (MTBITC) as
compared to the widely consumed conventional
variety, Aokubi (Nakamura et al. 2008). The
myrosinase, a cytosolic plant enzyme that
hydrolyses 4-methylthio-3-butenyl glucosinolate
(MTBGLS) into the natural pungent agent
4-methylthio-3-butenyl isothiocyanate (MTBITC),
was located exclusively in the outer epidermal
layer in Aokubi daikon, while 4-methylthio-3-bu-
tenyl glucosinolate (MTBGLS) was widely dis-
tributed throughout the root tissue. Despite the
skin being a potentially rich source of myrosinase
in Aokubi, the skin is usually peeled off in the cur-
rent practice of preparing daikon for cooking.
Thus, new practices were therefore proposed for
the preparation of daikon tubers that eliminated
the peeling of the skin to avoid removing the
enzyme needed to convert MTBGLS to the health-
beneficial MTBITC.
A total of eight glucosinolates were identified
in different tissues of radish, including five ali-
phatic glucosinolates (4-methylsulfinyl-3-butenyl
838
glucosinolate, 2-hydroxy-3-butenyl glucosino-
late, ethyl glucosinolate, 4-methylthio-3-butenyl
glucosinolate and 6-heptenyl glucosinolate) and
three indole glucosinolates (1-methoxyindol-
3-ylmethyl glucosinolate, indol-3-ylmethyl
glucosinolate, 4-hydroxyindol-3-ylmethyl gluco-
sinolate) (Li et al. 2008). The contents of total
glucosinolates in the edible roots and sprouts
were higher than that in the leaves. The major
glucosinolate in edible roots and sprouts was
4-methylthio-3-butenyl glucosinolate, account-
ing for 75.5 % and 71.5 % of the total glucosino-
lates, respectively. Indol-3-ylmethyl GS was the
major glucosinolate in radish leaves, accounting
for 57.1 % of the total leaf glucosinolates.
Other Phytochemicals
The contents of phenolic acids (hydroxycinnamic
acids and hydroxybenzoic acids) in radish roots
were much smaller than in the corresponding
leaves (Stöhr and Herrmann 1975). In small rad-
ishes, p-coumaric acid and, in radishes, ferulic
acid dominated after hydrolysis. Small radishes
showed higher concentrations of phenolic acids
in outer tissue layers. In contrary to hydroxycin-
namic acid derivatives, the contents of hydroxy-
benzoic acid derivatives (p-hydroxybenzoic,
vanillic, salicylic and gentisic acid) mostly
were small. Partially, hydroxycoumarins (aescu-
letin and scopoletin) were also identified.
N-methylphenethylamine, a secondary amine
with indirect sympathomimetic action (Marquardt
et al. 1976), and phenethylamine (Gutiérrez and
Perez 2004) were found in radish root.
Alkaloids, saponins and flavonoids and the fol-
lowing constituents were detected in the swollen
radish root: 11 long-chain alkyl esters, 3-methyl-
5-propylnonane, 5-(hydroxymethyl)undecane,
4,6-dimethyldodecane, n-tetradecane; n-hexadec-
ane, n-heneicosane; 2,6,10,14-tetramethylhexade
cane; n-eicosane, methyl n-tetradecanoate; 2,6,10,
15-tetramethylheptadecane; n-tricosane; n-hexa-
decanoic acid methyl ester; tritetracontane;
10-octadecenoic acid methyl ester; 9-octadece-
noic acid ethyl ester; 2 aromatic ketones, diphe-
nyl-benzophenone and 1-phenyl-pent-4-en-1-
one; a benzyl halide, a-iodotoluene; an aryl ether,
1-methoxy-4-(prop-2-en-1-yl)benzene; a quinone,
Brassicaceae
2,6-di-tert-butylquinone; an aromatic aldehyde,
2-benzylideneoctanal; a long-chain aromatic
alkane, 6-phenyldodecane; and a long-chain alkyl
nitrile, 9-octadecenitrile (San Jaun et al. 2012).
The enzyme cysteine synthase comprising of
two identical subunits of molecular weight
33,000 was isolated from radish roots (Tamura
et al. 1976). Radish root was found to contain
phospholipase D (Rakhimov et al. 1981).
Phospholipase D displayed a more pronounced
specificity for lecithin as substrate and was
weakly active in the reaction with cephalin. An
addition of Ca2+ increases the enzymatic affinity
for the substrate in all cases studied. Two cationic
isoperoxidases (designated C1 and C3) and four
anionic isoperoxidases (designated A1, A2, A3n
and A3) from Korean radish root were purified
(Lee and Kim 1994). All six isoperoxidases were
glycoproteins composed of a single polypeptide
chain. The MW of C1, C3, A1 and A2 were ca
44,000, while anionic isoperoxidases A3n and
A3 had MW of 31,000 and 50,000, respectively.
Deglycosylated A2 and C3 by trifluoromethane-
sulfonic acid treatment showed MWs of 37,000
and 40,000, respectively, suggesting that the car-
bohydrate contents for these isoenzymes were 14
and 9 %, respectively. Two chitin-binding pro-
teins with lysozyme activity, named as CBP1 and
CBP2, were purified to homogeneity with the
molecular weights of 26.9 kD and 24.8 kD,
respectively, from radish roots (Lai et al. 2006).
CBP1 and CBP2 were found to be bifunctional
enzymes with activities of lysozyme and chitin-
ase, but without chitosanase activity. A unique
acidic calcium-binding protein RVCaB, rich in
glutamic acid and proline and lacking aromatic
amino acid residues, was found in radish vacu-
oles and may be involved in the vacuole Ca(2+)
storage function (Ishijima et al. 2007). RVCaB
was found to be predominantly an unstructured
protein with a polyproline type II helix.
Arabinogalactan proteins (AGPs) were iso-
lated from primary and mature radish roots
(Tsumuraya et al. 1988). These root AGPs were
composed mainly of L-arabinose and D-galactose
but were distinguishable from each other in their
contents of L-fucose as well as of protein and
hydroxyproline. The structures of the carbohy-
Raphanus raphanistrum subsp. sativus
drate moieties of the root AGPs were essentially
similar to those of AGPs isolated from seeds and
mature leaves in that they consisted of consecu-
tive (1→3)-linked β-D-galactosyl backbone
chains having side chains of (1→6)-linked β-D-
galactosyl residues, to which α-L-arabinofuranosyl
residues were attached in the outer regions. Three
ferredoxin isoproteins (R-Fd A, R-Fd B-1 and
R-Fd B-2) were purified from white roots of
radish (Raphanus sativus L. var. acantiformis
cultivar Miyashige) (Wada et al. 1989).
1-O-sinapoyl-β-glucose:L-malate O-sinapoyl-
transferase (SMT) from cotyledons of red radish
was purified to apparent homogeneity with a
2,100-fold enrichment and a 4 % recovery
(Gräwe et al. 1992). Apparent molecular mass
of 52 and 51, respectively, were determined. On
isoelectric focusing, the SMT resolved into
two isoforms which showed slightly different
molecular mass (SMT I, Mr/isoelectric point =
51/5.75; SMT II, Mr/isoelectric point = 51.5/5.9).
The enzyme accepted all the hydroxycinnamic
acid glucose esters tested with relative ratios of
initial velocity values of 100∶85∶45∶26∶2.6
of 1-O-sinapoyl-, 1-O-feruloyl-, 1-O-caffeoyl-,
1,2-di-O-sinapoyl- and 1-O-(4-coumaroyl)-
β-glucose.
Genes involved in anthocyanin biosynthesis in
radish include phenylalanine ammonia lyase
(PAL), cinnamate 4-hydroxylase (C4H),
4-coumarate-CoA ligase (4CL), chalcone syn-
thase (CHS), chalcone isomerase (CHI), flava-
none 3-hydroxylase (F3H), dihydroflavonol
reductase (DFR) and anthocyanidin synthase
(ANS). RsDFR and RsANS were found to accu-
mulate in the flesh or skin of two radish cultivars
(Man Tang Hong and Hong Feng No. 1) (Park
et al. 2011). Radish skin contained higher CHS,
CHI and F3H transcript levels than radish flesh in
all three cultivars. In red radish, 16 anthocyanins
were separated and identified. Some of them
were acylated with coumaroyl, malonyl, feruloyl
and caffeoyl moieties. Furthermore, (-)-epicate-
chin and ferulic acid were also identified in the
three cultivars.
Myrosinase was found to accumulate in myro-
sin cells and in other non-specialised cells in the
seeds, seedling and other organs of radish plant
839
(Phelan et al. 1984). Two myrosinase isoenzymes
were extracted and purified from radish root tis-
sues with Mrs 28,800 and 58,900 (Jwanny et al.
1995). The most active one (120 U/mg) was iden-
tified and characterised. The yield of the purified
myrosinases was 21 mg (2,398 U) of pure
enzymes from 100 g of dry root tissues.
Myrosinase (thioglucoside glucohydrolase), a
plant enzyme that hydrolyses glucosinolates,
principally to isothiocyanates, was purified to
homogeneity in good yield from 8-day-old dai-
kon seedlings (Shikita et al. 1999). The purified
enzyme was resolved into two subunits with
molecular masses of 61 and 62 kDa. Ascorbic
acid activated the purified enzyme more than
100-fold. The enzyme also had β-glucosidase
activity and hydrolysed p-nitrophenyl-β-D-
glucopyranoside. Two myrosinase isoenzymes
were extracted and purified from radish root tis-
sues with Mrs 28,800 and 58,900 (Jwanny et al.
1995). The most active one (120 U/mg) was iden-
tified and characterised. The yield of the purified
myrosinases was 21 mg (2,398 U) of pure
enzymes from 100 g of dry root tissues. Two
cDNA clones of myrosinase (thioglucoside glu-
cohydrolase) were isolated from radish
(Raphanus sativus) seedlings (Hara et al. 2000).
Both clones were identified as MB (B type
myrosinase). The tissue distribution of gene
expression and enzyme activity of myrosinase
corresponded well to the site of glucosinolate
accumulation in different tissues of radish. The
myrosinase–glucosinolate system was localised
in the cotyledons in the seedlings and in the peel
of the root in the mature plant. Myrosinase
mRNA and activity were localised in the epider-
mis and vascular cambium that were present in
the peripheral part of the root, but few signals
were detected in the parenchyma inside of the
vascular cambium. Since the myrosinase–gluco-
sinolate system is known to be a defence system
in higher plants, the localisation of the myrosi-
nase–glucosinolate system in the peel of the
root may act to protect the sink organ from
the attack of herbivores or pathogens in soil.
In radish, the concentrations of individual
polycyclic aromatic hydrocarbons (PAHs) and
phthalic acid esters (PAEs) varied from
840
undetectable to 803 μg/kg dry weight (DW) and
from undetectable to 2,048 μgkg DW, respec-
tively (Cai et al. 2008). Compared to the control,
higher application rates of sewage sludge resulted
in pronounced increases in radish shoot, root and
soil concentrations of PAHs and PAEs. PAE con-
centrations in radish grown in latosolic red soil
spiked with sludge compost were higher, while
the PAH concentrations were comparable to
those receiving 10 g/kg of sewage sludge.
However, the root biomass of radish in soil
amended with compost was significantly higher,
and the shoot-to-root ratio was significantly
lower than in the other treatments. The biocon-
centration factors (BCFs, the ratio of contami-
nant concentration in plant tissue to the soil
concentration) of di-n-butyl phthalate and
di(2-ethylhexyl) phthalate in both shoots and
roots and of total PAH concentrations in roots
were less than 1.0, but some BCFs for individual
PAHs were high with a maximum value of 80.
Phytochemicals in Leaf/Stem
In radish leaves, the anaerobic accumulation
of alanine was accompanied by a loss of aspar-
tate, and these changes preceded gamma-
aminobutyrate accumulation and glutamate loss
(Streeter and Thompson 1972). Accumulation
of gamma-aminobutyrate was due to accelera-
tion of glutamate decarboxylation and arrest
of gamma-aminobutyrate transamination.
Changes in keto acid content did not appear to
be the cause of amino acid changes. Most of the
aspartate may be converted anaerobically to
alanine via oxaloacetate and pyruvate.
Regarding phenolic acids (hydroxycinnamic
acid compounds), leaves of radish (Raphanus
sativus var. sativus and var. niger) mainly con-
tained compounds of caffeic and p-coumaric
acid (Schmidtlein and Herrmann 1975). In the
group of hydroxybenzoic acid derivatives,
traces of salicylic and gentisic acid and fre-
quently vanillic acid were also found. Caffeic,
p-coumaric,, ferulic and sinapic acid esters of
malic acid and the enzyme(s) involved in their
syntheses were found in the vacuoles of radish
Brassicaceae
leaf protoplasts (Strack and Sharma 1985). The
flavonoid kaempferol 3,7-di-O-a-L-rhamnopy-
ranoside (lespedin) was identified in radish leaves
(Muminova et al. 2006). The following polyphe-
nolic compounds catechin, protocatechuic acid,
syringic acid, vanillic acid, ferulic acid, sinapic
acid, o-coumaric acid, myricetin, and quercetin
were found in radish leaves and stem (Beevi et al.
2010b). Oriental radish (Longipinnatus group)
was reported to contain the flavonol kaempferol
0.34 mg/100 g (Arai et al. 2000). Quercetin, rutin
and kaempferol were determined in radish leaves
by liquid chromatography–tandem mass spec-
trometry (Devaraj et al. 2011).
Radish leaf was characterised chiefly by
indole glucosinolates, largely 3-indolylmethyl-
glucosinolate (glucobrassicin) with traces of
4-methoxy-3-indolylmethyl-glucosinolate
(4-methoxyglucobrassicin) and 4-hydroxy-3-
indolylmethyl-glucosinolate (4-hydroxybrassicin)
(Sang et al. 1984). Hydroxylated cinnamic acid
malate esters, namely, 2-O-(p-coumaroyl)-L-
malate, 2-O-caffeoyl-L-malate and 2-O-feruloyl-
L-malate, were found to be quantitatively
predominating compounds in the fraction of car-
boxylic acids isolated from radish leaves and
inflorescences (Nielsen et al. 1984). The esters of
p-coumaric, ferulic and caffeic acid with malic
acid were isolated from the leaves, the ester of
sinapic acid with malic acid from cotyledons
of R. sativus (Brandl et al. 1984). The flavonoid
kaempferol 3,7-di-O-α-L-rhamnopyranoside (les-
pedin) was isolated from radish leaves (Muminova
et al. 2006).
The major volatile components found in the
oil of leaves of black, white and red radish in
respective sequence were phytol (65.3, 69.7,
65.9 %), hexadecanoic acid (palmitic acid)
(14.3, 2.5, 8.5 %), methyl linolenate (11.1, 8.9,
2.1 %) and (Z)-3-hexen-1-ol (0.7, 0.7, 6.9 %)
(Blažević and Mastelić 2009). Corresponding
aliphatic or aromatic glucosinolate compounds
identified in the leaves constituted only 0.3–
5.7 % of the isolated volatiles. They were
4-methylpentyl isothiocyanate (0, 0.1, 0 %),
5-(methylthio)-4-pentenenitrile (tr, 2.2, 0.3 %),
4-(methylthio)butyl isothiocyanate erucin
(0.3,1.7, tr %), 2-phenylethyl isothiocyanate (tr,
Raphanus raphanistrum subsp. sativus
0.4, 0 %), 4-(methylthio)-3-butenyl isothiocya-
nate (0, tr, 0 %), benzenepropanenitrile (0, tr,
0 %), 3-(methylthio)propyl isothiocyanate
(iberverin) (0, tr, 0 %), benzyl isothiocyanate (tr,
0, 0 %) and 2-phenylethyl isothiocyanate (tr, 0.4,
0 %). Alkanes identified in the leaves were
undecane (tr, 0.2, 0.2 %), dodecane (0.3, 1.1,
0.8 %), tridecane (0.1, 0.3, 0.3 %), tetradecane
(tr, tr, 0.1 %), tricosane (0.2, 0.3, 0.5 %) and tet-
racosane (0.1,0.2,0.2 %). Other aliphatic alco-
hols and carbonyls identified in the leaves were
(E)-2-hexenal (tr, tr, 0.8 %), nonanal (0, tr,
0.2 %), (E,E)-2,4-heptadienal (0.1,0, 0 %), benz-
aldehyde (tr, 0, 0 %), 2-phenylacetaldehyde (tr,
0.2, 0.2 %), 4-ethylbenzaldehyde (0, 0, 0.1 %),
isobutyl phthalate (0, 0.3, 0 %) and dibutyl
phthalate (0.4, 1.1, 1.0 %). Other fatty acids and
esters identified were tetradecanoic acid (myris-
tic acid) (tr, 0, 0 %) and methyl palmitate (0.1, 0,
0 %). Other sulfur and/or nitrogen compounds
identified were dimethyl trisulfide (tr, 0.4, 0.2 %)
and 1-(methylthio)-0-3-pentanone (0, tr, 1.1 %).
Besides phytol, terpenes identified were (E)-β-
ionone (0.5, 1.3, 1.4 %) and neophytadiene (1.5,
1.1, 1.3 %). Miscellaneous compound identified
included 4-vinyl-2-methoxyphenol (0.5, 1.3,
1.0 %), 2,3-dihydrobenzofuran (coumaran) (0.1,
tr, 0.4 %), β-cyclocitral (0, tr, tr), β-damascone
(0, 0.1, 0 %) and 1H-indole (0.1, 1.8, 0.2 %).
Nineteen volatile aglycones were identified for
the first time in the leaves of all three varieties of
radish. The main aglycones in the leaves of black,
white and red radish were, respectively, eugenol
(29.2, 30.2, 37.1 %), 2-phenylethanol (23.5, 2.0,
11.8 %), 4-vinyl-2-methoxyphenol (2.7, 12.4,
2.4 %), (Z)-3-hexen-1-ol (6.2, 2.6,10.4 %),
methyl vanillate (0, 10.0, 6.3 %), 1H-indole (6.8,
2.2, 0.7 %), benzyl alcohol (4.2, 1.4, 1.8 %), van-
illin (3.7, 3.5, 6.3 %) and methyl salicylate (3.4,
0.8, 0 %). Other aglycones included benzalde-
hyde (0.1, 1.4, 0 %), 2-phenylacetaldehyde (0,
0.9, 0 %), 1-phenylethyl alcohol (0.2, 0.1, 0.2 %),
2-methoxyphenol (guaiacol) (0.6, 1.7, 0 %),
2-methoxy-4-methylphenol (0, 2.2, 0 %), phenol
(0, 2.3, 0 %), 4-ethyl-2-methoxyphenol (2.6, 2.3,
0 %), 2,4-dimethylphenol (0, 3.2, 0.3 %), 4-keto-
α-ionol (0.5, 2.9, 2.1 %) and 3-hydroxy-β-ionone
(0, 2.4, 1.1 %).
841
Two isoproteins (L-Fd A and L-Fd B) were
isolated from radish leaves (Wada et al. 1989).
Three ferredoxin components, a, b and c, were
found in the green shoots of Japanese radish
seedlings and mature leaves (Obata et al. 1995).
Component a was further separated into two
components, a1 and a2. They found that the pri-
mary structures of components a1, a2 and b were
all found in 12 possible structures of L-Fd A, one
the two leaf ferredoxins, L-Fds A and B, isolated
by Wada et al. (1989). They concluded that L-Fd
A was a mixture of three ferredoxins correspond-
ing to components a1, a2 and b.
Two L-arabino-D-galactan-containing glyco-
proteins having a potent inhibitory activity
against eel anti-H agglutinin were isolated from
mature radish leaves and characterised to have a
similar monosaccharide composition that con-
sisted of L-arabinose, D-galactose, L-fucose,
4-O-methyl-D-glucuronic acid and D-glucuronic
acid residues (Tsumuraya et al. 1984). Two iso-
proteins (L-Fd A and L-Fd B) were isolated from
(Raphanus sativus L. var. acantiformis cultivar
Miyashige) leaves (Wada et al. 1989). Two
induced radish leaf proteins (designated Rs-AFP3
and Rs-AFP4) were purified and shown to be
homologous to seed Rs-AFPs and to exert similar
antifungal activity in-vitro (Terras et al. 1995). A
chimeric Rs-AFP2 gene under the control of the
constitutive cauliflower mosaic virus 35S pro-
moter conferred enhanced resistance to the foliar
pathogen Alternaria longipes in transgenic
tobacco. The term ‘plant defensins’ was proposed
to denote these defence-related proteins. On the
basis of molecular weight and their antifungal
properties, the protein samples from the
Alternaria alternata- and Fusarium oxysporum-
infected radish leaves were found to be PR2 (glu-
canase) and PR3 (chitinase) (Khanal et al. 2014).
Gibberellins (GA1, GA3, GA7) were reported
to play a direct role in the bolting of ‘Miyashige-
sofuto’ radish but were probably not directly
functional in initiating flowering (Suge and
Rappaport 1968). The concentrations of putative
active GAs (GA1 and GA4) and their precursors
(GA9 and GA20) in both the stems and leaves of
R. sativus grown without cold treatment were
higher in the long-day condition than the short-day
842
condition (Nakayama et al. 1995). The concen-
trations of the above four GAs in the stems and
leaves generally tended to decrease during the
cold treatment, although GA1 in the stems was at
almost the same level before and after the cold
treatment. These results suggested that the GA
biosynthesis was promoted by the long-day con-
dition rather than cold treatment. Considering
that application of GAs after the cold treatment
as well as cultivation in the long-day condition
after cold treatment induced the bolting of R.
sativus, both the cold treatment and activation of
GA biosynthesis by cultivation in the long-day
condition appeared to be essential to bolting. A
series of GAs belonging to either the early-13-
hydroxylation or early-non-hydroxylation path-
way were identified in four cultivars of Japanese
radish (Raphanus sativus L. cvs. Sofutori-
Miyashige, Wakayama, Hanashirazu-Hayafutori-
Tokinashi and Wase-Shijunichi), which varied
greatly in their cold requirement for bolting (or
stem elongation) and flowering, suggesting that
the endogenous active GAs were GA1 and GA4
(Nishijima et al. 1995). The concentration of GA1
increased in the stem and decreased in the leaf
during bolting in all the cultivars.
Two anti-migraine compounds were isolated
from radish roots and elucidated as cis-(1-
methylazetidin-2-yl)methanol and cyclo-
(4-methyl-val-4-methyl-val) (Wu et al. 2014).
Foliar applications of brassinosteroids (poly-
hydroxyl steroidal phytohormones), namely,
28-homobrassinolide and 24-epibrassinolide, to
the radish plants resulted in substantial increment
(65 %) in soluble protein contents and nucleic
acids DNA and RNA (Vardhini et al. 2012).
Increased levels of carbohydrates in terms of
reducing sugars, nonreducing sugars and starch
were also observed.
Seed Phytochemicals
The major active compounds in radish seeds
reported were alkaloids, glucosinolates, brassi-
nosteroids and flavonoids (Sham et al. 2013).
Two fructooligosaccharides, sucrose, glucose
and galactose were identified in radish and
Brassicaceae
Brassica napus seed ethanol extracts (Orlovskaya
et al. 2013).
The thiocyanate (SCN)− content in radish
seeds prior to planting was negatively correlated
with the SCN− content in marketable roots of
five white cultivars (Chong and Bible 1975). On
the contrary, there was a corresponding positive
correlation for eight red cultivars. Although
larger seeds resulted in larger radishes, there
was no difference in SCN− content in the root
of plants grown from different seed sizes.
Studies by Sang et al. ( 1984) found that the
only indole glucosinolate present in the rad-
ish seed meal was 4-hydroxy-3-indolylmethyl
GS (4-hydroxyglucobrassicin) with the dominant
glucosinolate being 4-methylsulfinylbut-3-enyl
GS (glucoraphenin), which contained an unsatu-
rated sulfinyl R group. Ten isothiocyanates, seven
aliphatic hydrocarbons and some other volatile
substances were characterised in Raphanus sati-
vus var. niger seeds (Afsharypuor and Balam
(2005). The main isothiocyanates were hexyl iso-
thiocyanate (18.4 %), 4-methylthiobutyl isothio-
cyanate (17 %), 4-methylpentyl isothiocyanate
(8.4 %), 4-methylthio-(3E)-butenyl isothiocya-
nate (5.2 %), 4-methylthio-(3Z)-butenyl isothio-
cyanate (4.7 %) and isoamyl isothiocyanate
(2.4 %). Seven 4-methylthio-butanyl derivatives
were identified in the radish seed methanol
extract: sinapoyl desulfoglucoraphenin, (E)-5-
(methylsulfinyl)pent-4-enoxylimidic acid methyl
ester, (S)-5-((methylsulfinyl)methyl)pyrrolidine-
2-thione, 5-(methylsulfinyl)-4-pentenenitrile,
5-(methylsulfinyl)-pentanenitrile, sulforaphene
and sulforaphane (Kim et al. 2014).
Radish seeds contained two major types of
storage protein aggregates which could be sepa-
rated by gel filtration into 12 and 1.7 Svedberg
fractions (Laroche et al. 1984). Radish 12
Svedberg particles comprised a series of nine
major polypeptides ranging from 33 to 30 kDa
and composed of six subunits approximately
55 kDa. Each subunit is a couple of two polypep-
tides linked by a disulfide bridge. The 1.7
Svedberg particle is composed of two polypep-
tides of 10 and 12 kDa and smaller peptides
of approximately 7 kDa. Twelve and 1.7
Svedberg particles also differed in their amino
Raphanus raphanistrum subsp. sativus
acid composition, the 1.7 Svedberg being
particularly rich in glutamic acid and proline. An
L-arabino-D-galactan and an L-arabino-D-galac-
tan-containing proteoglycan were isolated from
radish seeds (Tsumuraya et al. 1987). The proteo-
glycan consisted of 86 % of a polysaccharide
component containing β-L-arabinose and D-galac-
tose as major sugar constituents, together with
small proportions of D-xylose, D-glucose and
uronic acids and 9 % of a hydroxyproline-con-
taining protein. Raphanuside, an unusual
oxathiane-fused thioglycoside isolated from
Raphanus sativus seeds, was synthesised giving
11 % overall yield (Yang et al. 2010). Two novel
sulfur compounds S-6-(methylsulfinyl)methyl-
1,3-thiazinan-2-thione and O-ethyl N-(E)-4-
(methylsulfinyl)but-3-enylcarbamothioate were
isolated from the seeds (Zhang et al. 2010). A
novel compound rasatiol was isolated from rad-
ish seed used as Korean herbal medicine,
‘NaBokJa’ (Roh et al. 2013).
A basic β-galactosidase (β-Galase) with an
apparent molecular mass of 45 kDa was purified
281-fold from imbibed radish seeds (Sekimata
et al. 1989). The enzyme was maximally active at
pH 4.0 on p-nitrophenyl β-D-galactoside and
β-1,3-linked galactobiose. Radish seed and leaf
arabino-3,6-galactan proteins were resistant to
β-Galase alone but could be partially degraded by
the enzyme after the treatment with a fungal α-L-
arabinofuranosidase leaving some oligosaccha-
rides consisting of D-galactose, uronic acid,
L-arabinose and other minor sugar components
besides D-galactose as the main product. α-L-
arabinofuranosidase was purified 1,043-fold
from radish seeds (Hata et al. 1992). The purified
enzyme was a homogeneous glycoprotein con-
sisting of a single polypeptide with an apparent
molecular weight of 64,000 and an isoelectric
point value of 4.7. The enzyme characteristically
catalyses the hydrolysis of p-nitrophenyl
α-L-arabinofuranoside and p-nitrophenyl β-D-
xylopyranoside. The enzyme was shown to split
off α-L-arabinofuranosyl residues in sugar beet
arabinan, soybean arabinan-4-galactan and rad-
ish seed and leaf arabinogalactan proteins.
Arabinose and xylose were released by the action
of the enzyme on oat-spelt xylan. Synergistic
843
action of α-L-arabinofuranosidase and β-D-
galactosidase on radish seed arabinogalactan pro-
tein resulted in the extensive degradation of the
carbohydrate moiety. Radish seeds were found to
contain two homologous, small, 5 kD, cysteine-
rich oligomeric proteins designated Raphanus
sativus antifungal protein 1 (Rs-AFP1) and
Rs-AFP2; they were found in the cell wall occur-
ring predominantly in the outer cell layers lining
different seed organs (Terras et al. 1992a, 1995).
The radish 2S storage albumins in the seeds
were identified as the second novel class of
antifungal proteins. An α-L-arabinofuranosidase/
β-D-xylosidase was cloned from immature seeds
of radish by reverse transcriptase-PCR (Kotake
et al. 2006). It hydrolysed α-L-arabinofuranosyl
residues of the carbohydrate moieties of arabino-
galactan proteins (AGPs). The gene, designated
RsAraf1, was found to encode a bifunctional
α-L-arabinofuranosidase/β-D-xylosidase and the
results suggested that RsAraf1 was involved in
the hydrolysis of the carbohydrate moieties of
AGPs in immature radish seeds. Raphanus
sativus antifungal protein 1 (Rs-AFP1), a 51
amino acid residue plant defensin, was isolated
from radish seeds (Fant et al. 1998). Cationic
peroxidase Cs with molecular mass of 44 kDa
was purified to apparent homogeneity and
characterised from Korean radish seeds
(Kim and Lee 2005). The cationic peroxidase
Cs showed the peroxidase activities for native
substrates, such as coumaric acid, ferulic acid
and scopoletin. This result suggested that
cationic peroxidase Cs may play an important
role in plant cell wall formation during seed
germination. Rs-AFP2 (Raphanus sativus anti-
fungal peptide 2), an antifungal plant defensin,
was isolated from radish seed (Aerts et al. 2007).
Karri and Bharadwaja (2013) linked two plant
defensins, namely, Trigonella foenum-graecum
defensin 2 (Tfgd2) and Raphanus sativus anti-
fungal protein 2 (Rs-AFP2) genes, by a linker
peptide sequence (occurring in the seeds of
Impatiens balsamina) and made into a single-
fusion gene construct with enhanced antifungal
activity.
Eleven gibberellins (GAs) were identified in
extracts of mature seed of radish (Raphanus
844
sativus cv. Taibyosobutori) (Nakayama et al.
1990). These GAs comprised seven 13-hydroxy
GAs [GA1, 3-epi-GA1, GA8, GA17, GA19, GA20
and a new GA, 12α-hydroxy GA20 (GA77)] and
four non-13-hydroxy GAs [GA9, GA24,
12β-hydroxy GA24 (tentative) and GA25]. The
major GAs were GA8, GA20, GA24 and GA77.
During the first 48 hours after imbibition of
water by quiescent radish seeds, the changes in
ATP concentrations, oxygen utilisation and fresh
weights followed a triphasic time course, charac-
terised by a rapid initial increase, which extended
from 0 to approximately 1.5 hours, a lag phase
from 1.5 to 16 hours and a sharp linear increase
from 16 to 48 hours (Moreland et al. 1974). After
imbibition of water by the quiescent seeds, for
1 hour, the ATP concentration had increased to
2.5, and ADP and AMP concentrations had
decreased to 0.3 and 0.1 nmole/seed, respectively.
In unimbibed seeds, the concentrations of ATP,
ADP and AMP were <0.1, 0.9, and 2.2 nmoles/
seed, respectively. Oxidative phosphorylation
was estimated to have contributed 15, 20 and 65 %
of the pool ATP at 1.5, 16 and 48 hours, respectively.
Phytochemicals in Seedlings
and Sprouts
In dark-grown radish seedlings, the level of phe-
nylalanine transaminase was higher in cotyle-
dons than in the root and hypocotyls (Tomè et al.
1975). The maximum activity of phenylalanine
ammonia lyase (PAL) was found in the root.
Only PAL was significantly increased by light.
Continuous far-red light (active phytochrome P
fr) stimulated the synthesis of all prenyl chains
(C40 carotenoids; C45 in plastoquinone-9; C20
phytyl in chlorophylls, α-tocopherol and vitamin
K1) in plastids of etiolated radish seedlings but
had no or only little effect on the dark pattern of
the prenyl chain formation (Lichtenthaler and
Becker 1975). White light enhanced the accumu-
lation of prenyl chains to a much higher degree
than does far-red light. Two separate enzyme
activities for the formation of the 7- and
9-glucopyranosyl derivatives of the cytokinin
6-benzylaminopurine (BAP) were found in soluble
Brassicaceae
extracts of expanded cotyledons of radish and
purified (Entsch and Letham 1979). Each activity
resulted in the formation of both glucosides, and
the enzymes use UDP-glucose as the source of
the glucose residue.
Nucleoside polyphosphates, namely, nucleo-
side triphosphates, nucleoside tetraphosphates,
pentaphosphates and hexaphosphates were found
in labelled RNA preparations from radish seed-
lings (Aspart-Pascot et al. 1976). Polyadenylic
and polyadenylated ribonucleic acids were found
in radish seedlings (Aspart et al. 1979). Two life-
time classes of polyadenylic acid were deter-
mined in these seedlings: a short-lived component
with a half-life of 30 minutes which represented
60 % of poly(A) and a more stable component
with varying half-lives of which the majority
range from 4 to 10 hours and a few were consid-
erably longer. Anti-bolting activity was detected
in an extract of rosette shoots of radish plants by
an assay using seedlings cultured in-vitro (Yoshida
et al. 2010). The causal compound that strongly
inhibited bolting was isolated and identified as
α-(7Z,10Z,13Z)-hexadecatrienoic acid mono-
glyceride (16:3 monoglyceride). The compound
disappeared completely after vernalisation, and
bolting occurred thereafter. A membrane-bound
3-hydroxy-3-methylglutaryl-coenzyme A reduc-
tase was characterised from 4-day-old, dark-grown
radish seedlings (Bach et al. 1986).
The specificity of binding and the complete
precipitation of β-fructosidase activity by the
insolubilised lectin implied that all β-fructosidase
activity measured in Raphanus sativus seedling
extracts was linked to (a) glycoprotein form(s) of
this enzyme (Faye and Berjonneau 1979). When
36-hour-old dark-grown radish seedlings were
transferred to far-red light, there was a decrease
in glycoprotein cytoplasmic β-fructosidase
(betaF) and an increase in cell wall betaF com-
pared to the dark controls (Faye et al. 1986).
Cytoplasmic and cell wall-bound β-fructosidase
exhibited high antigenic similarities but differed
according to charge heterogeneity and carbohy-
drate microheterogeneity. Growth of radish seed-
lings in the presence of tunicamycin resulted in a
partial inhibition of betaF glycosylation, but non-
glycosylated betaF still accumulated in the cell
Raphanus raphanistrum subsp. sativus
wall under far-red light. The nonglycosylated
cytoplasmic and cell wall betaF forms have the
same relative molecular mass, but glycosylated
forms had different oligosaccharide side chains,
with respect to size and susceptibility to
α-mannosidase and endoglycosidase D digestion.
An inhibitor of Ca2+–calmodulin (Cam)-
dependent brain phosphodiesterase was present
in the soluble fraction of embryo axes from
ungerminated radish seeds (Cocucci and Negrini
1988). During the early phases of germination,
the fresh weight and the levels of DNA and RNA
of embryo axes increased, the level of the inhibi-
tor decreased and the level of Cam increased. It
was found that the Ca2+–calmodulin system was
activated in early germination of radish seeds by
an increase in Cam and a decrease in the inhibitor
levels; that fusicoccin, probably through the acti-
vation of membrane functions, increased Cam
level; and that the abscisic acid (ABA) inhibition
on germination was not mediated by the Ca2+–
Cam system. The peptide-N4-(N-acetyl-β-D-
glucosaminyl) asparagine amidase (PNGase)
activity was found in dry radish seeds, and its
level was constant during germination and post-
germination (Berger et al. 1995). The endo-N-
acetyl-beta-D-glucosaminidase (ENGase) activity
was first detected about 18 hours after the start of
imbibition (HAI) and displayed a maximum level
at 36 HAI. After 36 HAI, the production of both
enzymes was constant until days 4–5. Both
enzymes displayed substrate specificities corre-
sponding to the potential glycoprotein substrates
found in plants.
In radish hypocotyl, 4-hydroxy-3-indolylmethyl
glucosinolate (4-hydroxybrassicin), 4-methoxy-
3-indolylmethyl glucosinolate (4-methoxygluco-
brassicin) and 3-indolylmethyl glucosinolate
(glucobrassicin) were present with the major glu-
cosinolate being 4-methylthio-3-butenyl gluco-
sinolate (glucoraphasatin) (Sang et al. 1984). In
comparison to daikon seeds, glucoraphenin
(4-methylsulfinyl-3-butenyl Gs, GRE) content in
daikon sprouts decreased from about 90 to about
12 μmol/g of dry weight (DW), whereas a 25-fold
increase from about 3 to 76 μmol/g DW of the
glucoraphasatin (4-methylthio-3-butenyl Gs,
GRH) content was determined (Barillari et al.
845
2005). An efficient pure GRH gram-scale produc-
tion process from R. sativus (kaiware daikon)
sprouts resulted in significant yield improvement
of up to 2.2 % (DW basis). Radish sprouts con-
tained significantly greater concentrations of glu-
cosinolates (3.8-fold) and isothiocyanates
(8.2-fold) than the mature radish taproot and also
contained significantly greater concentrations of
phenolics on average 6.9-fold (Hanlon and Barnes
2011).
Malate, which plays many essential roles in
plant metabolism, was found to be a potent in-
vitro inhibitor of the cytosolic enzyme phospho-
enolpyruvate carboxylase in radish root hair
cytoplasm (Bodson et al. 1991). A novel α-L-
fucosyltransferase capable of transferring
L-fucose (L-Fuc) from GDP-L-Fuc to the O-2 of
α-L-arabinofuranosyl residue (GDP-L-Fuc–α-L-
arabinofuranoside 2-α-L-fucosyltransferase) was
found in the microsomal fraction of primary
roots from 6-day-old radish seedlings (Misawa
et al. 1996). Soluble and cell wall-bound gamma-
glutamyl transferases (GGTs) were purified
from radish cotyledons (Nakano et al. 2006).
Soluble GGTs (GGT I and II) had the same M(r)
of 63,000, and were composed of a heavy sub-
unit (M(r), 42,000) and a light one (M(r),
21,000). The properties of GGT I and II were
similar. Both soluble and bound GGTs utilised
glutathione, gamma-L-glutamyl-p-nitroanilide,
oxidised glutathione and the conjugate of gluta-
thione with monobromobimane as substrates and
were inhibited by acivicin, but soluble GGTs
were also distinguished from bound GGTs with
regard to these properties. Two forms of RD21
(responsive to desiccation-21), an Arabidopsis
cysteine protease, were identified from cotyle-
dons of daikon radish, consisting of an interme-
diate form (iRD21) containing a granulin domain
and a mature form (mRD21) lacking this domain
(Kikuchi et al. 2008). A cathepsin B-like cyste-
ine protease (CBCP) that was reported to play a
role in disease resistance and in protein remo-
bilisation during germination was purified from
daikon radish cotyledons (Tsuji et al. 2008). The
molecular mass of the enzyme was estimated to
be 28 kDa. The best synthetic substrate for
CBCP was t-butyloxycarbonyl Leu-Arg-Arg-4-
846
methylcoumaryl 7-amide. Daikon CBCP exhibited
both endopeptidase and exopeptidase activities.
In addition, CBCP was found to display car-
boxymonopeptidase activity against the sub-
strate o-aminobenzoyl-Phe-Arg-Phe(4-NO(2)).
Three acidic and three neutral growth inhibi-
tors were detected in Sakurajima radish hypocot-
yls grown in light (Hasegawa and Miyamoto
1978). Among them, all of the acidic and one of
the neutral inhibitors increased with the time
period of illumination, whereas the other two
neutral substances remained almost unchanged in
the light but decreased in the dark. Thus, the lev-
els of all six inhibitors were higher in light-grown
seedlings than in dark-grown ones. A neutral
growth inhibitor named raphanusol A was iso-
lated as a colourless powder from light-grown
radish seedlings and partially characterised as a
phenolic compound (Hasegawa and Miyamoto
1980). Raphanusol A, isolated from an acetone
extract of light-exposed seedlings of Sakurajima
radish, was characterised as 1-β,4-di-O-(4-
hydroxy-3,5-dimethoxycinnamoyl) gentiobiose
(Hase and Hasegawa 1982). Another growth
inhibitor, designated raphanusol B, was isolated
in crystalline form from light-grown Sakurajima
radish seedlings and has been shown to be
1-sinapoylglucose (Hasegawa and Hase 1981).
Raphanusol B inhibited the growth of intact and
excised hypocotyls of etiolated radish seedlings.
The raphanusol B content of the radish seedlings
increased greatly under red light but decreased in
the dark. Another neutral growth inhibitor, desig-
nated raphanusanin, was isolated in crystalline
form from light-exposed Sakurajima radish
(Raphanus sativus var. hortensis f. gigantissimus
Makino) seedlings and identified as a new com-
pound, 3-methoxy-4-methylthio-2-piperithione
(Hasegawa et al. 1982). Applied raphanusanin
inhibited the hypocotyl growth of etiolated radish
and lettuce seedlings at concentrations higher
than 1.5 × 10−6 M. A new growth inhibitor iso-
lated from an acetone extract of light-exposed
seedlings of Sakurajima radish was characterised
as 2-thioxothiazolidine-4-carboxylic acid (Hase
et al. 1983). Three growth inhibitors which might
be involved in phototropism of Sakurajima radish
(Raphanus sativus var. hortensis f. gigantissimus
Brassicaceae
Makino) hypocotyls were isolated as crystalline
forms from light-exposed radish seedlings and
identified as cis- and trans-raphanusanins and
6-methoxy-2,3,4,5-tetrahydro-1,3-oxazepin-2-
one (designated raphanusamide) (Hasegawa et al.
1986). The cis- and trans-raphanusanins inhib-
ited the growth of etiolated radish hypocotyls at
concentrations higher than 1.5 μmolar, rapha-
nusamide at concentrations higher than
20 μmolar. Studies found that cis- and trans-
raphanusanins caused growth inhibition by sup-
pressing the action of auxin and/or cytokinin
(Sakoda et al. 1991). A new growth inhibitor was
isolated from light-grown radish seedlings and its
structure determined as 3-(E)-(methylthio)
methylene-2-pyrrolidinethione (Sakoda et al.
1990). Above concentrations of 30 mg/L, it
inhibited hypocotyl growth in the etiolated cress
seedling test.
During radish germination, there was a wide
fluctuation of soluble sinapic acid esters. Two
major constituents were rapidly degraded in the
imbibition phase, and during cotyledon growth,
new soluble sinapic acid esters appeared, from
which three major components exhibited consid-
erable quantitative changes in opposing directions
(Strack 1977). Sinapine esterase that catalysed
sinapine (a major phenolic compound in radish
seeds) into sinapate and choline during seed ger-
mination was found in radish cotyledons
(Nurmann and Strack 1979). Sinapine was iso-
lated from radish seeds (Liu et al. 2002). The
mean recovery rate for sinapine thiocyanate from
radish seeds was 99.75 % and the amount ranged
from 1.696 to 16.96 (Li and Jing 2010). The fol-
lowing components were identified in the meta-
bolic pathway of sinapic acid and its ester
derivatives in radish cotyledons: sinapoylglucose,
sinapine (sinapoylcholine), sinapoylmalate and
disinapoylglucose (Linscheid et al. 1980). Protein
preparations from radish cotyledons converted the
sinapoyl moiety of 1-sinapoylglucose to L-malate
to form sinapoyl-L-malate via a sinapoyltransfer-
ase (Tkotz and Strack 1980). During the develop-
ment of radish seedlings, transconjugation
reactions of sinapine (O-sinapoylcholine) via
transiently accumulating the intermediate
1-O-sinapoyl-β-D-glucose in the cotyledons led to
Raphanus raphanistrum subsp. sativus
accumulation of O-sinapoyl-L-malate (Strack
1982). The first committed enzyme in sinapate
ester biosynthesis was established as a glucosyl-
transferase (UDP-glucose–sinapate glucosyl-
transferase) that transferred the glucose moiety
from UDP-glucose to the carboxyl group of
sinapate (Strack 1982). The resulting β-acetal
ester 1-O-sinapoyl-β-D-glucose (sinapoylglucose)
served as acyl donor in subsequent transacylation
reactions thus providing an alternative to CoA-
dependent pathways (Mock and Strack 1993). A
uridine 5′-diphosphoglucose–hydroxycinnamic
acid acyl-glucosyltransferase (HCA-GT) catalys-
ing the formation of 1-O-sinapoyl-β-glucose (EC
2.4.1.120) in seedlings of Raphanus sativus was
partially purified (Mock and Strack 1993). The
enzymes channelling sinapoylglucose into the
major accumulating sinapate ester compounds
had been defined as sinapoylglucose–malate
sinapoyltransferase (SMT) and sinapoylglucose–
choline sinapoyltransferase (SCT), respectively
(Tkotz and Strack 1980; Strack et al. 1986).
L-phenylalanine ammonia lyase (PAL) was not
involved in sinapoyl derivative metabolism in rad-
ish (Strack et al. 1978). In-vivo inhibition of PAL
activity with α-aminooxy-β-phenylpropionic acid
(AOP) did not affect the accumulation of sinap-
oylglucose, whereas the formation of anthocya-
nins, flavonols and a feruloyl derivative was
severely depressed. Administration of AOP to the
intact seedling revealed I50 values to be 0.14 mM
for pelargonidin, 0.16 mM for kaempferol and
0.18 mM for the feruloyl derivative. Biosynthesis
of phosphatidylcholine in young radish seedlings
was supplied with choline from degradation of the
seed constituent sinapine (sinapoylcholine)
(Strack 1981). During seedling development, the
quantity of labelled sinapine rapidly decreased as
a result of sinapine degradation with a concomi-
tant label increase in free choline, phosphorylcho-
line and phosphatidylcholine. Approximately
50 % of the choline liberated from sinapine was
consumed in the biosynthesis of phosphatidylcho-
line. Seedlings of red radish accumulated high
amounts of free malic acid and sinapoylmalate,
when grown on nitrate as the sole N-source
(Dahlbender and Strack 1984). In the presence of
ammonium (NO3−/NH4+-N, 1:2), both metabolites
847
failed to accumulate, and the levels of arginine,
asparagine, glutamine, histidine and serine were
greatly increased. Sinapic acid esters, namely,
1-sinapoylglucose, sinapoylmalate, 6,3′-disi-
napoylsucrose and 1,2-disinapoylglucose from
crude methanolic extracts from cotyledons of
6-day-old R. sativus seedlings, cultured on nutri-
ent solution with NO3−/NH4+-N, were higher than
when cultured on NO3−-N. The extractable activ-
ity of 1-sinapoylglucose–L-malate sinapoyltrans-
ferase, an enzyme which plays a key role in
channelling malic acid into the sinapic-acid
metabolism of this plant, was positively corre-
lated with the malic acid level in the cotyledons.
From the results, it was proposed that free malic
acid might be the likely candidate for regulating
the activity of 1-sinapoylglucose–L-malate
sinapoyltransferase. L-malate, sinapic acid esters
and 1-sinapoylglucose–L-malate sinapoyltrans-
ferase (SMT) catalysing the synthesis of sinapoyl-
L-malate were found in the vacuoles of protoplasts
obtained from cotyledons of red radish (Sharma
and Strack 1985). The epidermis and mesophyll
tissues of radish cotyledons were found to contain
sinapic acid esters (1-sinapoylglucose, sinapoyl-
L-malate, 6,3′-disinapoylsucrose), kaempferol
glycosides, free malic acid and enzyme involved
in the synthesis of sinapoyl-L-malate, 1-sinapoyl-
glucose–L-malate sinapoyltransferase (SMT)
(Strack et al. 1985). The kaempferol glycosides
were mainly localised in the upper epidermis of
radish seedling cotyledons, while the sinapoyl
esters were found in all tissues but differed mark-
edly in their concentrations. Disinapoylsucrose
was localised predominantly in the mesophyll,
while most sinapoylmalate was found in the epi-
dermal layers, as was most 1-sinapoylglucose–
L-malate sinapoyltransferase (SMT) activity.
Epidermal sinapoyl esters were restricted to guard
cells and to guard mother cells and adjacent
epidermal cells. Strack et al. (1986) afforded
evidence indicating that the 1-sinapolyglucose–
L-malate sinapoyltransferase (SMT) activity in
radish cotyledons might be related to the metabo-
lism of malic acid.
Sinapoyl glucose was found in radish
sprouts (Herrmann 1989). Methyl sinapate,
1,2-disinapoyl-β-D-glucopyranoside and β-D-(3,4-
848
disinapoyl)fructofuranosyl-α- D -(6- sinapoyl)-
glucopyranoside were isolated from radish sprouts
(Takaya et al. 2003). The main phenolic acids rep-
resented in R. sativus grown in hydroponic culture
in excess copper were chlorogenic, vanillic, caf-
feic, syringic, p-coumaric and ferulic acids,
whereas the least represented were gallic, proto-
catechuic and p-hydroxybenzoic acids (Sgherri
et al. 2003). The Cu contents increased with the
treatment in both shoots and roots, maintaining in
the roots a value eight- to tenfold higher than in
the shoots. In both parts, phytochelatin-SH con-
tent reached the maximum at 5 μM copper and
then decreased, reaching at 15 μM copper the
control value in the roots and a value fivefold
higher than the control value in the shoots. The
phenolic acids as well as the total and reduced
ascorbate contents increased with the intensifica-
tion of copper treatment. Notwithstanding these
changes, total ascorbate remained 35 % higher in
the shoots than in the roots.
Phytochemicals in Flowers
Radish stigma diffusates were found to contain
many protein bands; the molecular weights of
some of the major fractions were estimated to be
15,000, 30,000–46,000 and 70,000 Da (Zhang
et al. 1983). Glycine, glutamic acid, serine and
aspartic acid were some of the predominant
amino acids. The carbohydrate fraction of the
glycoprotein consisted of arabinose 17.3 %,
galactose 19.1 %, xylose 8.1 %, mannose 5.4 %,
glucose 23.7 %, rhamnose and/or fucose 26.4 %.
In the stigma surface diffusates, the content of
protein was estimated to be 16 % and that carbo-
hydrate was 11 %.
Antioxidant/Redox Activity
The antioxidative activity of hot water extract of
daikon was higher than that of the ambient water
extract (Katsuzaki et al. 2004). One of the anti-
oxidants was isolated as L-tryptophan which
changed to 5-hydroxytryptophan, another anti-
oxidant in the rat liver microsome. Radish root
Brassicaceae
extract reduced the levels of thiobarbituric acid
reactive substance significantly in all treated
albino rats as compared to the experimental con-
trol group (Chaturvedi 2008). It also increased
the levels of reduced glutathione and increased
the activity of catalase. The extract also inhibited
in-vitro cumene hydroperoxide-induced lipid
peroxidation. The results suggested that radish
root extract afforded protection by strengthening
the antioxidants like glutathione and catalase.
Among the different extraction solvents,
methanolic extract of the leaves and stem showed
potent reductive capacity, significantly inhibited
linoleic acid peroxidation and displayed metal
chelating activity (Beevi et al. 2010b).
Additionally, they scavenged free radicals effec-
tively with IC50 of 31 and 42 μg/mL for DPPH
radical, 23 and 52 μg/mL for superoxide radical,
67 and 197 μg/mL for hydrogen peroxide and 56
and 62 μg/mL for nitric oxide, respectively. The
leaves showed the most potent antioxidant and
radical scavenging activity as compared to the
stem, attributable to the higher polyphenolic con-
tent. The methanolic radish root extract showed
significant ferric reducing ability, moderate metal
chelating activity and strong radical scavenging
activity (Beevi et al. 2012). Polyphenolic content
in radish was estimated to be in the range 13.18–
63.54 mg/g dry weight, with a considerable
amount being obtained with polar solvents.
Catechin was found to be the most abundant phe-
nolic compound in water extract and sinapic acid,
the predominant phenolic compound in methano-
lic, ethyl acetate and hexane extracts.
Red radish extract in which the major com-
pounds were acylated pelargonidin derivatives
appeared to form a complex with Fe3+ or Cu2+
(Wang et al. 2010). It displayed a concentration-
dependant reducing power and scavenging effect
against 2,2′-azino-bis-(3-ethylbenzothiazoline-
6-sulfonic acid) radicals (with IC50 = 1.74 mM).
It could promote the cleavage of plasmid DNA
with Cu(II)/H(2)O(2) or Cu(II) alone. The
extract also showed growth inhibition of Bel-
7402 cells at lower concentration. The results
suggested that the formation of reactive oxygen
species might be involved in the mechanism of
DNA damage. The acylated pelargonidin deriva-
Raphanus raphanistrum subsp. sativus
tives extracted from red radish could act as anti-
oxidant and pro-oxidant, and their antioxidant
and pro-oxidant properties were relative to the
reaction conditions.
Hydrogen peroxide oxidation of glucorapha-
satin (4-methylthio-3-butenyl GS) readily resulted
in complete transformation into glucoraphenin
(4-methylsulfinyl-3-butenyl GS). ABTS*+ radi-
cal caused complete decay of the glucosinolate in
radish sprouts (Barillari et al. 2005). Even though
not directly related to its radical scavenging
activity, the assessed reducing capacity of gluco-
raphenin suggests that R. sativus sprouts might
possess potential for health benefits. Spraying
exogenous plant hormone methyl jasmonate
(MeJA) upon radish sprout significantly increased
the total phenolic content that resulted in the
increased DPPH* (2,2-diphenyl-1-picrylhydrazyl)
free radical scavenging capacity (Kim et al.
2006). In addition, the phenylalanine ammonia
lyase activity also increased by 60 % at 24 hours
after MeJA treatment. However, the same
treatment decreased the amount of 4-methylthio-
3-butenyl isothiocyanate (MTBITC), a major
isothiocyanate in radish sprout, and the activity of
myrosinase, an enzyme related to produce
isothiocyanates.
Glucoraphasatin (GRH) and its corresponding
isothiocyanate (ITC), 4-methylthio-3-butenyl
isothiocyanate (GRH-ITC), from radish sprouts,
were able to quench the 2,2-diphenyl-1-
picrylhydrazyl (DPPH) radical, with second-
order rate constants of 14.0 and 43.1 M−1 s−1,
respectively (at 298 K in methanol), whereas the
corresponding value measured for the reference
antioxidant α-tocopherol was 425 M−1 s−1 (Papi
et al. 2008). GRH reacted with H2O2 and tert-
butyl hydroperoxide in water (pH 7.4) at
37 °C, with rate constants of 1.9 × 10−2 and
9.5 × 10−4 M−1 s−1 (paralleling recently developed
synthetic antioxidants) being quantitatively
(>97 %) converted to glucoraphenin.
Radish sprouts were between 9- and 59-fold
more potent than the corresponding mature tap-
root at activating the antioxidant response ele-
ment (ARE) in a stably transfected hepatoma cell
line (Hanlon and Barnes 2011). The ARE activity
849
of the radish sprouts and mature taproots was
significantly correlated with the total isothiocya-
nate concentration of the radishes.
Among the 11 kinds of commonly available
vegetables, the methanol extract of radish sprout
(Japanese name ‘kaiware daikon’) exhibited the
highest hydroxyl radical scavenging potency
using the bleomycin–Fe method (1.8 times as
L-ascorbic acid) (Takaya et al. 2003). Several
sinapinic acid esters and flavonoids were isolated
with high radical scavenging potency, which may
contribute substantially to the activity. Black rad-
ish root juice exhibited significant antioxidant
properties (Lugasi et al. 2005). Supplementation
of the lipid-rich diet (20 % sunflower oil, 2 %
cholesterol, 0.5 % cholic acid in normal chow)
with black radish juice resulted in a significant
improvement of antioxidant enzyme activities
and the free radical scavenging capacity in hyper-
lipidaemic rats.
Anticancer Activity
The methanol extract of radish sprouts induced
quinone reductase activity in a dose-dependent
manner in the concentration range of 0.2–1.6 mg/
mL with a maximum of a 3.5-fold increase in
induction in murine Hepa1c1c7 cells (Lee and
Lee 2006). The induction of quinone reductase
by the extract was regulated at the transcriptional
level. The dichloromethane (CH2Cl2) fraction of
the extract showed the highest induction potency,
while the other fractions were less potent. The
results indicated that radish sprouts could be
regarded as a safe and promising new dietary
source for decreasing the risk of developing can-
cer as quinone reductase is known to play critical
roles in protection against chemical carcinogens
and other toxic xenobiotics.
Isothiocyanates from radish sprouts, 4-
methylthio-3-butenyl isothiocyanate (GRH-ITC)
and 4-methylsulfinyl-3-butenyl isothiocyanate
(GRE-ITC), reduced cell proliferation in a
dose-dependent manner and induced apoptosis
in three human colon carcinoma cell lines
(LoVo, HCT-116 and HT-29) (Papi et al. 2008).
850
The compounds significantly increased Bax and
decreased Bcl-2 protein expression, as well as
produce caspase-9 and PARP-1 cleavage after
3 days of exposure in the three cancer cell lines.
GRH-ITC treatment was shown to have no toxic-
ity with regard to normal human lymphocytes
(−15 %) in comparison with sulforaphane from
broccoli (complete growth inhibition). GRH and
GRH-ITC were able to quench the 2,2-diphenyl-
1-picrylhydrazyl radical. Barillari et al. (2008)
found that administration of kaiware daikon
extract (KDE), in combination with myrosinase
at doses corresponding to 50 μM 4-methylthio-3-
butenyl isothiocyanate (GRH-ITC) plus 15 μM
4-methylsulfinyl-3-butenyl isothiocyanate (GRE-
ITC) (50 μM KDE-ITC) to three human cancer
cell lines (LoVo, HCT-116 and HT-29), signifi-
cantly reduced cell growth by 94–96 % of control
in 6 days, outperforming pure GRH-ITC or GRE-
ITC at the same dose. In contrast, the same treat-
ment had no significant toxicity on normal human
T lymphocytes. A 50 μM concentration of KDE-
ITC had relevant apoptosis induction in all tested
cancer cell lines. Unlike pure GRH or GRH-ITC,
KDE also had significant chain-breaking antioxi-
dant activity, retarding the 2,2′-azobis-(2-
amidinopropane hydrochloride) (AAPH)-initiated
autoxidation of methyl linoleate in sodium
dodecyl sulfate (SDS) micelles at concentrations
as low as 4.4 ppm (−50 % in oxygen consump-
tion rate), and induced very fast quenching of
DPPH radical. In another study, Kim et al. (2011)
reported that the ethanol extract of radish leaves
inhibited cell proliferation of MDA-MB-231
human breast cancer cells after 48 hours of incu-
bation via the ErbB–Akt pathway. The extract
significantly decreased protein expression and
mRNA of ErbB-3 (epidermal growth factor
receptor) and mRNA expression of Akt (protein
kinase B); it increased significantly protein and
mRNA expression of Bax and Bcl-2 (apoptosis
regulators). Seven 4-methylthio-butanyl deriva-
tives from radish seeds showed antiproliferative
activity against the HCT-15 cancer cell, with IC50
values of 8.49–23.97 μM (Kim et al. 2014).
Nitrogen mustards cis-1-methoxy-2-deoxy-2-
[N,N-bis(2 -chloroethyl)amino]spirobrassinol
and trans-1-methoxy-2-deoxy-2-[N,N-bis(2 -chlo-
Brassicaceae
roethyl)amino]spirobrassinol derived from
1-methoxyspirobrassinol, an indole phytoalexin
produced by the Japanese radish, were designed
as prospective dual-action compounds with DNA-
alkylating effect and glutathione-depleting effects
that may sensitise cancer cells to alkylating
agents (Mezencev et al. 2009). Both new com-
pounds demonstrated cytostatic/cytotoxic effects
on various leukaemia and ovarian cancer cell
lines and dsDNA-destabilising effects in-vitro.
The cis isomer was the more promising of the
two compounds, exerting earlier onset of anticancer
effects on Jurkat cells via induction of apoptosis
compared to the traditional alkylating anticancer
agent melphalan. Additionally, it demonstrated
higher potency on ovarian cancer OVCAR3 cell
line and lower fold resistance between Jurkat and
Jurkat-M cells selected for the resistance to mel-
phalan. Mustard oil in ‘Shibori Daikon’ a variety
of Japanese radish selectively inhibited the pro-
liferation of H-ras-transformed derivative of rat
fibroblast HR-3Y1-2, but not 3Y1 rat fibroblasts
after 24 hours of treatment (Yamasaki et al.
2009). The selective inhibition was associated
with transient oxidative stress via reduced gluta-
thione (GSH) depletion and cell cycle G2/M
arrest. The mustard oil extract was found to contain
95.6 % of 4-methylthio-3-butenyl isothiocyanate
and 4.4 % of 4-methylthiobutyl isothiocyanate.
Hexane extract of radish root inhibited cell
proliferation and induced apoptosis in human
cancer cells by upregulation of pro-apoptotic
genes and downregulation of antiapoptotic genes
along with activation of Caspase-3 (Beevi et al.
2010a). The presence of several isothiocyanates
(ITCs) such as 4-(methylthio)-3-butenyl isothio-
cyanate (MTBITC), 4-(methylthio)-3-butyl
isothiocyanate (erucin), 4-methylpentyl isothio-
cyanate, 4-pentenyl isothiocyanate and sul-
foraphene was detected in the hexane extract.
4-(methylthio)-3-butenyl isothiocyanate (MTBITC)
extracted from Raphanus sativus significantly
inhibited nitroblue tetrazolium reduction by
superoxide radicals in a nonenzymatic superox-
ide generating system, to scavenge free radicals
and to cause a decrease in murine leukaemia cell
line (L1210) cell growth (Salah-Abbès et al. 2010b).
It also counteracted zearalenone oxidative stress
Raphanus raphanistrum subsp. sativus
to BALB/c mice keratinocyte cell line (C5-O)
through caspase-8 inhibition of apoptosis.
A low transformation of selenium into organic
forms was observed in radish plants grown in
Se(VI)-enriched culture media (Pedrero et al.
2006). On the contrary, in those plants exposed
to selenite, >95 % of the total selenium was
found as selenocysteine (SeCys2), selenomethi-
onine (SeMet) and Se-methylselenocysteine
(SeMetSeCys). The concentrations of these
compounds in fresh samples remained almost
unaltered after a simulated gastrointestinal
digestion. Therefore, a high selenium content of
Se-methylselenocysteine (65 %), previously
reported as a cancer chemopreventive species,
remained in the potentially bioabsorbable frac-
tion. The results suggested that radish enriched
in selenite could be a good choice as an organose-
lenium supplement for the human diet and
animal feed.
Antimutagenic Activity
A correlation was found between the potency
of antimutagenicity and the amount of 4-
(methylthio)-3-butenyl isothiocyanate (MTBITC)
in the n-hexane extracts of eight strains of daikon
(Nakamura et al. 2001). Because the pure MTBITC
also showed antimutagenicity, MTBITC was pre-
sumed to be the active antimutagen principle in
n-hexane extracts of daikon. In addition,
phenethyl isothiocyanate was found in a lesser
amount (5–33 nmol/100 g) in eight strains of dai-
kon, and allyl isothiocyanate and benzyl isothio-
cyanate were not detectable in any strains
(<3 nmol/100 g). The amount of total isothiocya-
nate in grated daikon was 7.0 times higher than
that in cut daikon measured after 30 minutes
of cooking. Through eating habits, humans
might be able to consume substantial amounts of
the antimutagen MTBITC from dishes using the
grated form of wild strains of daikon. 4-(methylthio)-
3-butenyl isothiocyanate (MTBITC) extracted
from Tunisian radish was found to prevent
genotoxicity and clastogenicity of zearalenone
in BALB/c mice and in in-vitro (Salah-Abbès
et al. 2009c).
851
Antidiabetic Activity
Normal rats fed with a diet containing Japanese
radish sprout (JRS) had lower plasma levels of
total cholesterol (TC), triglycerides (TG), phos-
pholipids (PL), fructosamine, glucose and insulin
and higher plasma levels of low-density lipopro-
tein cholesterol, whereas the JRS-fed
streptozotocin-induced diabetic rats showed
lower plasma levels of fructosamine, glucose and
insulin without changes in the plasma lipid
parameters (Taniguchi et al. 2006). JRS also
decreased the hepatic TC, TG and PL levels in
the normal rats and the TG level in the diabetic
rats. The results showed that JRS had a hypogly-
caemic activity in both the normal and diabetic
rats and partly improved lipid metabolism in the
normal rats. JRS also had the potential to allevi-
ate hyperglycaemia in cases of diabetes and may
serve in the primary prevention of diabetes mel-
litus. In subsequent study, they found that the
water-soluble (WSE) and fat-soluble extracts
(FSE) from Japanese radish sprouts exerted dier-
ring effects on carbohydrate and lipid metabo-
lism in normal and streptozotocin-induced
diabetic rats (Taniguchi et al. 2007). FSE sup-
pressed insulin secretion and improved lipid
metabolism in the normal rats. The effect of WSE
was different from that of the FSE as it decreased
blood glucose levels without increasing insulin
secretion and also lowered glycoalbumin and
fructosamine levels in the streptozotocin (STZ)-
induced diabetic rats.
Radish root juice extract at a dose of 300 mg/
kg BW was identified as the most effective dose
lowering blood glucose level (BGL) by 33.4 %
at 6 hours during fasting blood glucose studies in
normal rats (Shukla et al. 2011a). However, the
glucose tolerance test (GTT) revealed the maxi-
mum reduction of 15.9 % in BGL at 3 hours in
normal rats with the same dose, whereas the
reduction observed was by 23.8 and 28.3 % in
streptozotocin (STZ)-induced sub- and mild-
diabetic rats, respectively, at the same interval of
time. The data suggested that radish root juice
had hypoglycaemic potential coupled with anti-
diabetic efficacy.
852
Anti-inflammatory Activity
The 4-methylthio-butanyl derivative, sinapoyl
desulfoglucoraphenin, isolated from radish seeds,
exhibited anti-neuroinflammatory effect in
lipopolysaccharide-stimulated murine microglia
BV-2 cells (Kim et al. 2014). It significantly
inhibited nitrite oxide production with IC50 val-
ues of 45.36 μM. Moreover, it also reduced the
protein expression of inducible nitric oxide
synthase.
After treatment with granules from black rad-
ish root, all of the histopathological changes and
parameters of the redox state (epithelial lining
disruption, reduction in the number of entero-
cytes and goblet cells and the presence of inflam-
matory cells) caused by the fat-rich diet were
improved (Sipos et al. 2002). The structure of the
epithelial cells was similar to the controls, the
number of goblet cells increased and no inflam-
mation was observed.
Both radish leaf juice and root juice signifi-
cantly reduced carrageenan- and formalin-
induced paw oedema in rats, but radish leaf juice
compared to root juice produced more significant
anti-inflammatory effect in both acute and
chronic models of inflammation (Kamble et al.
2013). However, the anti-inflammatory effect of
radish leaf juice was less than the standard drug
diclofenac sodium.
Spasmogenic Activity
Aqueous extract of radish seeds containing ter-
penes, flavonoids, phenols, alkaloids and sapo-
nins showed a spasmogenic effect in isolated
rabbit jejunum and ileum, rat stomach fundus and
ileum and guinea pig ileum and jejunum (Ghayur
et al. 2005). The extract was around ten times
more potent in the guinea pig tissues, and this
effect was resistant to atropine, pyrilamine or
SB203186, while the spasmogenic effect in the
rat and rabbit tissues was atropine sensitive. The
extract exhibited atropine-sensitive GI (gastroin-
testinal) prokinetic and laxative effects in-vivo in
mice. In the atropinised rabbit jejunum, radish
seed extract produced a spasmolytic effect inde-
pendent of Ca2+ or K+ channels and adrenergic or
Brassicaceae
opioid receptor involvement. Activity-directed
fractionation of the extract yielded four fractions,
all showing effects similar to that of the parent
extract. The extract and its fractions were found
to be nonlethal up to 10 g/kg in mice for 24 hours,
except for the petroleum fraction, which showed
50 % mortality at high doses. Some known radish
compounds (spermine, spermidine, putrescine
and sinigrin) were also tested and found to be
devoid of any activity. A mild relaxant effect was
also observed in rabbit jejunum at the lower
doses (0.1–0.3 mg/mL) but not against K+-
induced contractions, ruling out a calcium
channel-blocking effect. In guinea pig ileum, rad-
ish leaf extract exhibited a stimulant effect resis-
tant to atropine while sensitive to pyrilamine
pretreatment. The findings suggested the pres-
ence of species-dependent gastrointestinal effects
of radish mediated partially through cholinergic
receptors in rabbit and rat tissues but through his-
taminergic activation in the guinea pig. The crude
radish leaf extract exhibited a dose-dependent
(0.03–5.0 mg/mL) spasmogenicity in guinea pig
ileum and colon (Gilani and Ghayur 2004). The
effect was insensitive to atropine pretreatment
but was completely abolished by pyrilamine indi-
cating involvement of histaminergic (H(1))
receptors. The contractile effect at high doses
(3.0–5.0 mg/mL) was followed by relaxation.
The extract also enhanced the transit of charcoal
meal in mice at 30–100 mg/kg. The petroleum
spirit, chloroform and aqueous fractions all
showed histaminergic activity in the ileum, aque-
ous fraction being more potent. The results indi-
cated the presence of a histaminergic
component(s) along with a weak spasmolytic
factor thus providing sound mechanistic basis for
the traditional use of the plant in constipation. In
subsequent studies, they found that administra-
tion of radish and betel nut extracts to isolated
rabbit gall bladder tissues modulated gall bladder
contractility in a concentration-dependent man-
ner similar to carbachol, a muscarinic receptor
agonist (Ghayur and Gilani 2012). The stimulant
effect of the extract, as well as that of carbachol,
was completely blocked in the presence of atro-
pine, a muscarinic antagonist, indicating similar-
ity in the mechanism of action of the extracts
with carbachol. The result showed the potential
Raphanus raphanistrum subsp. sativus
of these extracts to contract the gall bladder and
to subsequently increase bile secretion.
Hepatoprotective Activity
Acute and chronic administration of radish leaf
powder, its water and ethanol extracts signifi-
cantly decreased the elevated activity of serum
glutamate oxaloacetate transaminase (SGOT),
serum glutamate aspartate transaminase (SGPT),
serum lactate dehydrogenase (SLDH), serum
alkaline phosphatase (SAP) and serum total bili-
rubin in paracetamol-induced hepatotoxicity in
rabbits (Anwar and Ahmad 2006). Crude radish
leaf powder, its water and ethanol extracts pro-
duced nonsignificant effect on total protein con-
tents. In a separate study, administration of radish
root methanol extract to albino rats exerted a pro-
tective effect on paracetamol-induced hepatotox-
icity in a dose-dependent manner (Chaturvedi
et al. 2007; Chaturdevi and Machacha 2007).
Radish extract reduced the levels of thiobarbitu-
ric acid reactive substances (TBARS), SGOT and
SGPT and increased the level of GSH and
reduced glutathione (GSH) and catalase. The
extract reduced lipid peroxidation induced by
paracetamol and reverted the levels of SGOT and
SGPT to normal, indicating liver recovery. It also
restored GSH levels and recovery of catalase
activity. Oral administration of radish extract of
200 and 400 mg/kg body weight protected Wistar
albino rats against carbon tetrachloride-induced
hepatotoxicity (Mohammed et al. 2008). The
extract inhibited the increase in serum aspartate
aminotransferase (AST), alanine aminotransfer-
ase (ALT), alkaline phosphatase (ALP) and bili-
rubin concentrations besides adverse
histopathological changes induced by CCL4.
Studies found that oral administration of the
sulfur-radish extract and of sulforaphane after
CCL4-induced liver injury in mice both decreased
the serum level of alanine aminotransferase,
reduced the necrotic zones, inhibited lipid per-
oxidation and induced phase 2 enzymes without
affecting cytochrome P450 2E1 (CYP2E1) (Baek
et al. 2008). The results suggested that the admin-
istration of the sulfur-radish extract and of sul-
foraphane may partially prevent CCL4-induced
853
hepatotoxicity, possibly by indirectly acting as an
antioxidant by improving the detoxification
system.
Administration of radish extract enhanced the
antioxidant status and protected against oxidative
stress in the liver and kidney induced by zearale-
none (a nonsteroidal oestrogenic mycotoxin) in
BALB/c mice (Salah-Abbès et al. 2008b).
Co-treatment of male BALB/c mice with R. sati-
vus extract plus zearalenone prevented the hepa-
totoxic damage effects of zearalenone
(Salah-Abbès et al. 2009b). Radish extract suc-
ceeded in reversing hepatotoxic condition back to
normal levels for all studied parameters.
Radish enzyme extract showed hepatoprotec-
tive activity on tacrine-induced cytotoxicity in
HepG2 cells with EC50 value of 1,250 μg/mL
(Lee et al. 2012). Oral administration of the
extract at doses of 50 and 100 mg/kg and silyma-
rin at a dose of 50 mg/kg significantly reduced
the elevated levels of serum enzyme markers
induced by CCL4.
Enzyme Detoxification Activity
The crude aqueous extract from 0.3 to 3 mg of dry
Spanish black radish material (SBR) increased
the activity of the phase II detoxification enzyme
quinone reductase in the human hepatoma HepG2
cell line with a maximal effect at a concentration
of 1 mg/mL (Hanlon et al. 2007). Treatment of
HepG2 cells with the crude aqueous extract of
1 mg of SBR per mL also significantly induced
the expression of mRNA corresponding to the
phase I detoxification enzymes, cytochrome P450
(CYP)1A1, CYP1A2 and CYP1B1, as well as the
phase II detoxification enzymes, quinone reduc-
tase, haem oxygenase 1 and thioredoxin reductase
1. They showed that while glucoraphasatin addi-
tion was ineffective, the isothiocyanate metabolite
of glucoraphasatin, 4-methylthio-3-butenyl
isothiocyanate (MIBITC), significantly induced
phase II detoxification enzymes at a concentration
of 10 μM. Further, Hanlon et al. (2009) demon-
strated that the crude aqueous extract of Spanish
black radish and the isothiocyanate metabolite of
glucoraphasatin, 4-methylthio-3-butenyl isothio-
cyanate (MIBITC), were potent inducers of phase
854
I detoxification enzymes, cytochrome P450
(CYP)1A1, CYP1A2 and CYP1B1, as well as the
phase II detoxification enzymes, quinone reduc-
tase, haem oxygenase 1 and thioredoxin reductase
1 in the HepG2 cell line.
In HepG2 cells, raphasatin induced quinone
reductase activity and the RNA expression of
several phase 1 and 2 detoxification enzymes by
a significantly greater amount than the degrada-
tion products of raphasatin (Scholl et al. 2011).
Raphasatin, but not its degradation products, acti-
vated the antioxidant response element (ARE) in
a stably transfected reporter cell line. Mice fed a
diet consisting of 20 % freeze-dried radishes for
2 weeks had significantly higher liver expression
of cytochrome P450 (CYP)1A1, 1A2, quinone
reductase, microsomal epoxide hydrolase and
glutathione S-transferase α2 than mice fed with a
nutritionally matched control diet.
In an animal study, N’jai et al. (2012) found
expression of phase I and II detoxification enzymes
was significantly greater for mice fed with Spanish
black radishes than control diet. Six hours after
7,12-dimethylbenz(a)anthracene (DMBA) admin-
istration, the blood levels of DMBA in mice fed
with the radish diet were significantly lower than
mice fed with a control diet. DMBA reduced bone
marrow cells in mice fed with a control diet to a
significantly greater extent than mice fed with the
black radish diet. Colony-forming assays demon-
strated that mice on black radish diet had (1) less
reduction in lymphoid CFU-preB progenitor cells,
(2) greater recovery of CFU-preB progenitor cells
at 168 hours and (3) less reduction of CFU-GM
progenitor cells at 6 hours. Thus, mice fed with a
20 % black radish diet for 2 weeks had greater
expression of detoxification enzymes, faster
metabolism of DMBA and a reduction in DMBA-
induced bone marrow toxicity.
Antihypertensive Activity
Radish seed extract tested positive for the pres-
ence of saponins, flavonoids, tannins, phenols
and alkaloids and caused a dose-dependent
(0.1–3 mg/kg) fall in blood pressure and heart
rate of rats that was mediated via an atropine-
Brassicaceae
sensitive pathway (Ghayur and Gilani 2006). In
isolated guinea pig atria, the extract showed
dose-dependent (0.03–3.0 mg/mL) inhibition of
force and rate of contractions. In the atropine-
treated tissues, the inhibitory effect was abol-
ished and a cardiac stimulant effect was unmasked
which was resistant to adrenergic and serotoner-
gic receptor blockade. In the endothelium-intact
rat aorta, the extract inhibited phenylephrine-
induced contractions, which was blocked by atro-
pine and Nomega-Nitro-L-arginine methyl ester
hydrochloride which was also absent in the
endothelium-denuded preparations. The data
showed that the cardiovascular inhibitory effects
of the plant were mediated through activation of
muscarinic receptors thus possibly justifying its
use in hypertension. Administration of soluble
alkaloids of Raphani Semen significantly low-
ered the blood pressure of spontaneously hyper-
tensive rats (SHR) and improved the process of
cardiovascular remodelling (Li et al. 2007). Laiju
extract (LJE) from Raphani Semen and Flos
Chrysanthemi exhibited antihypertensive effect
in renal hypertensive rat (RHR) and spontaneous
hypertensive rat (SHR) (Chen et al. 2007).
Compared with saline control, blood pressure
was significantly lowered at 6 and 5 hours in high
and moderate LJE, respectively, in both RHR and
SHR groups. However, blood pressure was sig-
nificantly lowered at 2 and 3 hours in low LJE
in both RHR and SHR groups, respectively.
Compared with saline control, blood pressure
remained significantly lower in SHR in all dos-
age groups with a single daily dose for 28 days
of study. The results suggested that LJE had
potential in the preventive management of
hypertension.
The flavonoid kaempferol 3,7-di-O-α-L-
rhamnopyranoside (lespedin) isolated from rad-
ish leaves was found to have hypotensor activity
(Muminova et al. 2006).
Antiatherosclerotic Activity
The abnormal growth of vascular smooth mus-
cle cells (VSMC) is a prominent feature of vas-
cular disease, including atherosclerosis and
Raphanus raphanistrum subsp. sativus
restenosis after angioplasty. Treatment with
Korean white radish extract decreased the via-
bility of VSMC by 35 % after 24 hours treatment
(Suh et al. 2006). Radish extract showed potent
inhibitory effects on the DNA synthesis of cul-
tured VSMC. In addition, radish extract induced
apoptosis using cell death ELISA assay. These
inhibitory effects were associated with G1 cell
cycle arrest. Four active isothiocyanate (ITC)
were isolated from the hexane radish
extract 4-(methylthio)-3-butenyl isothiocyanate
(MTBITC), allyl isothiocyanate (AITC), benzyl
isothiocyanate (BITC) and phenethyl isothiocy-
anate (PEITC). When the VSMC were treated
with ITC, the cell viability was significantly
decreased.
Antimicrobial Activity
An antibacterial principle designated raphanin
was isolated from radish seeds (Ivanovics and
Horvath 1947). It was inhibitory to the growth of
Staphylococcus and coliform bacteria. The crude
juices of Raphanus sativus was found to be
strongly inhibitory in-vitro against Escherichia
coli, Pseudomonas pyocyaneus, Salmonella
typhi and Bacillus subtilis (Abdou et al. 1972).
An antifungal nonspecific lipid transfer protein
composed of two 9 KD subunits and possessing
43 amino acids was isolated from radish seed
(Terras et al. 1992b). In low ionic strength
medium, it exhibited in-vitro antifungal activity
with IC50 values of 48 μg/mL for Alternaria
brassicicola, 41 μg/mL for Alternaria pisi,
45 μg/mL for Botrytis cinerea, 25 μg/mL for
Colletotrichum lindemuthianum, 20 μg/mL for
Fusarium culmorum, 54 μg/mL for Fusarium
oxysporum f.sp. lycopersici, 58 μg/mL for
Fusarium oxysporum f.sp. pisi, 100 μg/mL for
Nectria haematococca, 18 μg/mL for Phoma
betae, 10 μg/mL for Pyricularia oryzae, 30 μg/
mL for Trichoderma hamatum and 7 μg/mL for
Verticillium dahliae. Two homologous, 5 kD,
cysteine-rich proteins designated Raphanus sati-
vus antifungal protein 1 (Rs-AFP1) and 2
(Rs-AFP2) from radish seeds were found to
exhibit potent antifungal activity in-vitro (Terras
855
et al. 1992b, 1995). Their antibiotic activity
showed a high degree of specificity to filamentous
fungi. The radish 2S storage albumins in the
seeds were identified as the second novel class of
antifungal proteins. All isoforms inhibit the
growth of different plant pathogenic fungi and
some bacteria. However, their antimicrobial
activities were strongly antagonised by cations.
Two induced radish leaf proteins (designated
Rs-AFP3 and Rs-AFP4) were purified and
shown to be homologous to seed Rs-AFPs and to
exert similar antifungal activity in-vitro (Terras
et al. 1995). Rs-AFP2, a 51 amino acid cysteine-
rich peptide isolated from radish seeds, exhib-
ited potent inhibitory activity against filamentous
fungi (Alves et al. 1994). A cDNA clone encod-
ing the Rs-AFP2 preprotein was modified by
recombinant DNA methods to allow expression
in the yeast Saccharomyces cerevisiae.
4-methylthio-3-butenyl isothiocyanate
(MTBI), the pungent principle in radish, exhib-
ited antimicrobial activity in-vitro (Uda et al.
1993b). Among the tested Gram-negative bacte-
rial strains (Escherichia coli, Enterobacter cloa-
cae, Salmonella typhimurium, Proteus vulgaris),
the growth of E. cloacae was most strongly
affected; the growth inhibition of the other
Gram-negative bacterial strains was relatively
lower. In contrast, the Gram-positive strains
(Staphylococcus aureus S-6, Staphylococcus epi-
dermidis, Bacillus subtilis, Bacillus cereus var.
mycoides) were more sensitive to 2.5–7.5 μmol
MTBI. Growth of B. cereus was highly inhibited,
and S. aureus was also relatively sensitive. At
2.5–7.5 μmol, MTBI inhibited the growth of the
yeasts (Candida valida, Debaryomyces hansenii,
Hansenula anomala) and fungi (Alternaria
helianthi, Cladosporium colocasiae, Eurotium
chevalieri, Penicillium frequentans, Mucor race-
mosus f. racemosus) far more strongly than that
of the bacteria. Especially, the growth of
A. helianthi and C. colocasiae was almost com-
pletely inhibited by 5.0–7.5 μmol of MTBI. A
strong inhibitory effect was also observed in
E. chevalier and the yeast C. valida. Two antimi-
crobial components were found in the water-
soluble products obtained by degradation
of 4-methylthio-3-butenyl isothiocyanate, the
856
pungent principle of radish, of which the major
one was isolated and indentified to be 2-thioxo-3-
pyrrolidinecarbaldehyde (TPC) (Uda et al.
1993a). The identified compound exhibited
prominent growth inhibition on the fungi
(Alternaria helianthi, Cladosporium colocasiae,
Eurotium chevalieri, Mucor racemosus f. race-
mosus, Penicillium frequentans, Penicillium
expansum, Penicillium martensii, Aspergillus
candidus, Aspergillus fumigatus) and Gram-
positive bacteria (Staphylococcus aureus S-6,
Staphylococcus epidermidis, Bacillus subtilis,
Bacillus cereus var. mycoides, Streptococcus fae-
calis). The inhibitory effect on the yeasts
(Candida valida, Candida lactis-condensi,
Debaryomyces hansenii, Hansenula anomala,
Zygosaccharomyces rouxii) and Gram-negative
bacteria (Enterobacter cloacae, Salmonella
typhimurium, Proteus vulgaris, Pseudomonas
aeruginosa) except for Escherichia coli was the
least among the tested microorganisms. The min-
imum inhibitory concentration (MIC) of
2-thioxo-3-pyrrolidinecarbaldehyde (TPC) was
determined by strains of microbes (Matsuoka
et al. 1997). MICs of 100 μg/mL were obtained
for the fungi Alternaria helianthi, Aspergillus
candidus and Aspergillus fumigatus; MICs of
200 μg/mL for Cladosporium colocasiae,
Eurotium chevalieri, Neurospora crassa; and
MICs of 400 ug/mL for Mucor racemosus f. rac-
emosus, Penicillium frequentans, Penicillium
expansum and Penicillium martensii. Yeasts were
less sensitive with MICs of 400 μg/mL for
Candida albicans and Saccharomyces bayanus
and 800 μg/mL for Candida valida, Debaryomyces
hansenii and Hansenula anomala. MICs obtained
for Gram-negative bacteria were 200 μg/mL for
Enterobacter cloacae, Pseudomonas aeruginosa
and Salmonella typhimurium and 100 μg/mL for
Escherichia coli and Proteus vulgaris. For the
Gram-positive bacteria, the most sensitive was
Bacillus subtilis with MIC of 50 μg/mL, followed
by Staphylococcus aureus and Staphylococcus
epidermidis at 100 μg/mL and Bacillus cereus
and Enterococcus faecalis at 200 μg/mL. The
antifungal and antibacterial actions of TPC were
due to the sporicidal and bactericidal activities.
Brassicaceae
In S. epidermidis, a dose-dependent inhibition of
the uptakes of both oxygen and radioactive pre-
cursors was observed, suggesting that TPC
caused damage to the mitochondrial functions
and biosynthetic systems. The ethanol and meth-
anol extracts of radish seeds exhibited in-vitro
antibacterial activity against Escherichia coli,
Klebsiella pneumonia, Proteus vulgaris,
Pseudomonas aeruginosa, Staphylococcus
aureus, Shigella sonnei, Salmonella typhi and
Salmonella paratyphi (Ahmad et al. 2013).
Rs-AFP2 (Raphanus sativus antifungal pep-
tide 2), an antifungal plant defensin isolated from
radish seed, was found to interact with glucosyl-
ceramides (GlcCer) in membranes of susceptible
yeast and fungi and induced membrane permeabi-
lisation and fungal cell death (Aerts et al. 2007). It
was shown that Rs-AFP2 induced reactive oxy-
gen species (ROS) in Candida albicans wild type
in a dose-dependent manner, but not at all in an
Rs-AFP2-resistant Deltagcs C. albicans mutant
that lacked the Rs-AFP2-binding site in its mem-
branes. The findings indicated that upstream bind-
ing of Rs-AFP2 to GlcCer was essential for ROS
production leading to yeast cell death. They also
demonstrated that deletion of C. albicans meta-
caspase 1, encoding the only reported (putative)
caspase in C. albicans, significantly affected cas-
pase activation by the apoptotic stimulus acetic
acid, but not by Rs-AFP2 (Aerts et al. 2009).
Their data indicated the existence of at least two
different types of caspases or caspase-like prote-
ases in C. albicans.
The acetone and hexane fractions of radish
root, stem and leaf exhibited selective antibacte-
rial activity against Bacillus subtilis,
Staphylococcus aureus, Staphylococcus epider-
midis, Enterococcus faecalis, Salmonella
typhimurium, Enterobacter aerogenes,
Enterobacter cloacae and Escherichia coli
(Beevi et al. 2009). Antibacterial activity was
strongest in the acetone fraction of root with
larger zone of inhibition and lower minimum
inhibitory concentration. The results obtained
were comparable to that with standard antibiot-
ics. Of the different parts of R. sativus studied,
root tended to be more active than the stem and
Raphanus raphanistrum subsp. sativus
leaf extracts in inhibiting the bacterial growth.
Five different ITCs such as allyl isothiocyanate
(AITC), phenyl isothiocyanate (PITC), benzyl
isothiocyanate (BITC), phenethyl isothiocya-
nate and 4-(methylthio)-3-butenyl isothiocya-
nate (MTBITC) were identified in different
parts of the plant. The low linear correlation
between the total ITC content and antibacterial
activity implied that bacterial growth inhibitory
ability of R. sativus was not dependent on the
total ITC content. However, the antibacterial
activity of R. sativus was well correlated with
AITC, PITC and BITC for all organisms except
for Enterococcus faecalis, whose inhibitory effect
was more related to MTBITC.
Radish root juice exhibited considerable
antimicrobial activity in-vitro against five bacte-
rial strains, viz., Klebsiella pneumoniae,
Staphylococcus aureus, Pseudomonas aerugi-
nosa, Enterococcus faecalis and Escherichia
coli with minimum inhibitory concentration
(MIC) ranging from 0.078 to 0.625 mg/mL
(Shukla et al. 2011b). Ethanolic and ethyl acetate
extracts of radish root peels were the most effec-
tive of all extracts against bacterial strains tested
(Janjua et al. 2013). Ethyl acetate extract of rad-
ish root peels was most effective against
Staphylococcus aureus, Bacillus subtilis,
Salmonella typhi and Klebsiella pneumonia. The
ethanol extract had the highest zone of inhibition
against Micrococcus luteus, Pseudomonas aeru-
ginosa, Bordetella bronchiseptica and
Enterobacter aerogenes.
Antiviral Activity
Administration of black radish aqueous extract
by intranasal instillations to mice before inocula-
tion of the influenza virus A/PR 8/34 (H1N1)
strain protected against the experimental influ-
enza infection (Prahoveanu and Eşanu 1987,
1990). A significant decrease of the haemaggluti-
nin titre of the mouse lung homogenate was
noted, as well as a decrease of the mortality rate
and a significant increase of the rate of survival as
compared to the untreated control.
857
Antilithiatic and Diuretic Activities
Administration of the aqueous extract of radish
bark to rats with artificially induced urolithiasis
significantly decreased the weight of stones com-
pared to untreated urolithiatic rats (Vargas et al.
1999). This extract showed an increase in the
24 hour urine volume as compared to the control.
After treatment with black radish root juice for
6 days of female C57BL/6 mice with induced
gall bladder lithiasis, cholesterol gallstones were
eradicated significantly in the gall bladder of
mice; cholesterol and triglycerides levels were
decreased, and there was also an increase in
levels of HDL (Castro-Torres et al. 2012, 2014).
Immunoprotective Activity
Administration of radish extract was effective in
protecting against zearalenone-induced immuno-
logical disorders in BALB/c mice (Salah-Abbès
et al. 2008a). Mice treated with radish extract (5,
10 or 15 mg/kg) for 7 days before, during or after
zearalenone treatment showed a significant
improvement in lymphocyte, immunoglobulin
profile, T-cell subtypes, B cells and proinflamma-
tory cytokines. Moreover, treatment with the
highest dose of radish extract (15 mg/k) enhanced
the release of tumour necrosis factor-alpha and
interleukin 1beta, but the other parameters were
comparable with those of the control. Radish
extract was found to be effective for the protec-
tion of high-dose zearalenone immunotoxication
in BALB/c mice (Salah-Abbès et al. 2010a). The
extract at 15 and 30 mg/kg BW reduced the del-
eterious effects in immunological parameters of
high subchronic doses of 40 and 80 mg of zeara-
lenone/kg BW on modulation of lipopolysaccharide
(LPS). Radish extract was found to be effective
for the protection of high-dose zearalenone
immunotoxication in BALB/c mice (Salah-
Abbès et al. 2010a). The extract at 15 and 30 mg/
kg BW reduced the deleterious effects in immu-
nological parameters of high subchronic doses of
40 and 80 mg of zearalenone/kg BW on modula-
tion of lipopolysaccharide (LPS).
858
Salah-Abbès et al. (2014) found that radish
extract prevented cadmium-induced immuno-
toxic and biochemical alterations in rats.
Treatment with CdCl2 alone resulted in signifi-
cant decreases in plasma levels of total protein,
triglycerides, creatine kinase, creatinine, IgG
and IgA, T-lymphocyte subtypes (CD4+, CD3+,
CD56+ and CD8+) and thymic and hepatic indi-
ces (relative weights). Conversely, CdCl2 treat-
ment caused significant increases in serum LDH,
AST and ALT, in the formation/release of proin-
flammatory cytokines (IL-1 and TNF-α) and in
the relative weights of host spleen and kidneys.
Rats treated with radish extract alone had no dis-
cernable changes compared to the controls with
regard to all test parameters. Combined treat-
ment of CdCl2 and radish extract at any dose
resulted in a significant improvement of all test
parameters compared to those seen with cad-
mium alone.
Choleretic Activity
Studies showed that administration of 1.5 g/kg of
body weight radish sprouts (Kaiware Daikon)
extract (containing 10.5 % w/w glucosinolate
glucoraphasatin) for four consecutive days had
antioxidant properties and significantly induced
bile flow in rats (Barillari et al. 2006).
Gastroprotective Activity
Oral administration of freshly squeezed radish
juice (FRJ) in doses of 2 and 4 mL/200 g BW sig-
nificantly inhibited gastric ulcer formation
induced by necrotising agents (ethanol, sodium
hydroxide and sodium chloride), hypothermic
restraint stress and indomethacin (Algasoumi
et al. 2008). The FRJ also replenished the ethanol-
induced depleted gastric wall mucus secretion
and nonprotein sulfhydryl (NPSH) concentra-
tions in rats. Phytochemical screening showed
the presence of flavonoids, anthocyanins and sul-
furated constituents.
Brassicaceae
Studies by Ahn et al. (2013) found that pre-
treatment with purple Bordeaux radish (PBR)
extract at doses of 500 and 1,000 mg/kg, but not
250 mg/kg, significantly ameliorated ethanol-
induced gastric haemorrhages in rats. The immu-
noreactivities of inducible nitric oxide (iNOS)
and its by-product nitrotyrosine in the gastric
mucosa of ethanol-treated rats pretreated with
500 mg/kg PBR extract were significantly
reduced, as compared with rats treated with etha-
nol alone, the findings suggested that the gastro-
protective effect of PBR could be mediated partly
by the antioxidative activity of the extract.
Antidiarrhoeal Activity
Sinapine at 300 and 600 mg/Kg reduced the fre-
quency or incidence of purging induced by castor
oil or senna in mice, in a dose-dependent manner
(Zhang and Shen 1996). Sinapine inhibited the
gastrointestinal propellant rate of charcoal ink in
normal mice, but the inhibition was weak and
was not enhanced in a dose-dependent manner.
Skin Anti-ageing/Whitening Activity
The freeze-dried radish root juice showed higher
potency of tyrosinase inhibition (IC50 = 3.09 mg/
mL) than the methanolic extract (IC50 = 9.62 mg/
mL) (Jakmatakul et al. 2009). Also, the scaveng-
ing effects of the freeze-dried juice on DPPH
radical, superoxide anion radical and singlet oxy-
gen were greater than the methanolic extract,
with the respective IC50 values of 0.64, 4.20 and
1.42 mg/mL for the freeze-dried juice and 1.25,
6.28 and 2.40 mg/mL for the methanolic extract.
The contents of total phenolics, total flavonoids
and L-ascorbic acid (as per 1 mg of the dried
extract) were found to be 10.09, 0.51 and
24.11 μg for the freeze-dried juice and 6.59, 0.33
and 8.28 μg for the methanolic extract, respec-
tively. The higher contents of phenolic com-
pounds and L-ascorbic acid in the freeze-dried
juice appeared to be responsible for its greater
Raphanus raphanistrum subsp. sativus
antityrosinase and antioxidant activities. The
extract of R. sativus root appeared to be a good
candidate for application as a natural skin whit-
ening/skin anti-ageing agent due to its abilities to
inhibit tyrosinase and scavenge several types of
reactive oxygen species.
Rasatiol, isolated from radish seed, acceler-
ated dermal fibroblast growth in a dose-dependent
manner and increased the production of type 1
collagen, fibronectin and elastin (Roh et al.
2013). Phosphorylation of p42/44 extracellular
signal-regulated kinase, p38 mitogen-activated
protein kinase and Akt was remarkably increased
by rasatiol, indicating that enhanced extracellular
matrix production was linked to the activation of
intracellular signalling cascades. The results
indicated that rasatiol stimulated the fibrous com-
ponents of production and may be applied to the
maintenance of skin texture.
Reproductive Toxicity Protective
Activity
Radish extract, rich in many antioxidant com-
pounds, was safe and succeeded in counteracting
the oxidative stress and protected against
zearalenone-induced toxicological effects
(decreased sperm number, testosterone level and
antioxidant enzyme status) in male BALB/c mice
(Salah-Abbès et al. 2009a). The extract also
exhibited antigenotoxic effect in germ cells.
Laxative Activity
Administration of radish leaf aqueous extract and
radish leaf juice to Wistar rats exerted signifi-
cantly increased faecal output in loperamide-
induced constipation and laxative activity test
(Dande et al. 2014). Both the extract and the juice
increased the distance covered in charcoal meal
test and increased the water–ion secretion in elec-
trolyte secretion test indicating laxative activity
of Raphanus sativus leaf.
859
Antithyroid Activity
Makhkamov and Latipov (1965) reported a
decrease in thyroid hormone in rats fed with rad-
ish. Upon hydrolysis, 3-indolylmethyl glucosino-
late was reported to produce the thiocyanate
(SCN) ion which, when administered over a pro-
longed period, will reduce the iodine available to
the thyroid for incorporation into the thyroid hor-
mone (Greer 1950).
Boiled extracts of radish showed maximum
inhibition of thyroid peroxidase activity in-vitro
followed by cooked and raw extracts (Chandra
et al. 2004). After chronic radish feeding,
increased weight of thyroid gland, decreased thy-
roid peroxidase activity, reduced thyroid hor-
mone profiles and elevated level of thyrotropin
were observed in albino rats resembling a relative
state of hypoactive thyroid gland in comparison
to control even after supplementation of adequate
iodine (Chandra et al. 2006).
Food Safety Studies
Biogenic amines putrescine, cadaverine, histamine,
tyramine, spermidine and spermine increased dur-
ing broccoli and radish sprout production although
these levels were below those permitted by legisla-
tion (5 mg/100 g of edible food) (Martínez-
Villaluenga et al. 2008). Broccoli and radish
sprouts contained numbers of mesophilic, psychro-
trophic, total and faecal coliform bacteria which
were the usual counts for minimally processed ger-
minated seeds. They found that broccoli and radish
sprouts demonstrated no toxic effects on prolifera-
tion and viability of HL-60 cells and should be
included in our diets as healthy and safe fresh
foods. Raphani Semen processed by roasting was
reported to exhibit some adverse effects on mice
(Sham et al. 2013). Additionally, erucic acid, the
main fatty acid in Raphani Semen, was shown to
enhance the toxicity of doxorubicin. Thus, Raphani
Semen has a potential risk of causing toxicity and
drug interaction.
860
Allergy Issues
A case of allergic contact dermatitis from the rad-
ish was reported in a waitress aged 38 years
(Mitchell and Jordan 1974). The mixture of sini-
grin, the thioglucoside of allyl isothiocyanate,
mixed in petrolatum with the enzyme myrosinase
produced a positive patch test reaction. A more
recent case of generalised urticaria after inges-
tion of radish was reported by Damiani et al.
2011).
Traditional Medicinal Uses
Almost all parts of radish plant including leaves,
seeds and roots are utilised in medicine (Mayer
1981; Stuart 2013). The fresh juice obtained from
leaves is diuretic, laxative, and used for diarrhoea,
dropsy and general anasarca. Roots are considered
stimulant, carminative and corrective, antiscorbu-
tic and used for urinary complaints, haemorrhoids,
gastrodynia pains and various gastric ailments.
Roots are crushed and applied locally as dressing
or poultice for burns, scalds, ecchymoses or foetid
or smelly feet. Root decoction is employed for
fever and to bring out the rash in eruptive fevers.
Seeds are considered expectorant, digestive,
diuretic, laxative, stimulant, carminative and litho-
triptic and used to promote the flow of urine, bowel
movements and menstruation. Flowers are consid-
ered cholagogue.
Lai fu zi (radish) is employed in traditional
Chinese medicine (TCM) for treatment of gastro-
intestinal disorder and as expectorant in China
(Zhang et al. 2010). Radish is a preventive and
cure for stone (in the bladder or kidney) (Abdou
et al. 1972). It cures readily bronchial troubles
and whooping cough (so common among infants
in the East). Radish is reputed to be highly
diuretic. Radish is a cruciferous vegetable that
has been traditionally used in South Asia for dif-
ferent gastrointestinal, gall bladder and hepatic
diseases (Ghayur and Gilani 2012). In Mexico,
black radish is employed for the treatment of
gallstones and for decreasing lipid serum levels
(Castro-Torres et al. 2012, 2014).
Brassicaceae
Dried ripe radish seed (Raphani Semen) is
listed in Pharmacopoeia of the People’s Republic
of China to be commonly used in TCM for pro-
moting digestion, relieving distension, directing
‘Qi’ downwards and dissipating phlegm (Chinese
Pharmacopoeia Commission 2010; Tan et al.
2005). Raphani Semen is traditionally used to
treat food dyspeptic retention, distending pain in
the epigastrium and abdomen, constipation, diar-
rhoea and dysentery, panting and cough with
phlegm congestion clinically in combination
with other TCM herbs. For example, Raphani
Semen is one of the three important ingredients
of San-Zi-Yang-Qin-Tang, which is a common
TCM formula for relieving cough and asthma,
dissipating phlegm and promoting digestion.
Radish root has been used as a traditional anti-
migraine drug in China for hundreds of years
(Wu et al. 2014).
Other Uses
Raphanus sativus root was found to have poten-
tial in removal of phenolic compounds from
phenol-contaminated water (Naghibi et al. 2003).
Sliced radish or its juice was added separately as
enzyme source to phenol solution, and after
3 hours, more than 90 % of phenol were removed
in both cases.
Domingos et al. (2008) employed response
surface methodology (RSM) to determine the
optimum condition for the ethanolysis of R. sati-
vus crude oil. Three process variables were eval-
uated at two levels (2(3) experimental design):
the ethanol–oil molar ratio (6:1 and 12:1), the
catalyst concentration in relation to oil mass (0.4
and 0.8 wt.% NaOH) and the alcoholysis tem-
perature (45 and 65 °C). Radish press cake, a
solid residue from biodiesel processing, could be
used to produce adsorbents by microwave ther-
mal activation for the removal of cationic dyes
from wastewaters (Nunes et al. 2011). Both the
removal efficiency and the removal capacity
decreased with an increase in temperature, point-
ing towards the exothermic nature of the removal
process.
Raphanus raphanistrum subsp. sativus
Karri and Bharadwaja (2013) linked two plant
defensins, namely, Trigonella foenum-graecum
defensin 2 (Tfgd2) and Raphanus sativus anti-
fungal protein 2 (Rs-AFP2) genes, by a linker
peptide sequence (occurring in the seeds of
Impatiens balsamina) and made into a single-
fusion gene construct. This method produced
biologically active recombinant His6-tagged
fusion protein, which exhibited potent antifungal
action towards the plant pathogenic fungi
(Botrytis cinerea, Fusarium moniliforme,
Fusarium oxysporum, Phaeoisariopsis personata
and Rhizoctonia solani along with an oomycete
pathogen Phytophthora parasitica var. nicoti-
anae) at lower concentrations under in-vitro con-
ditions. This strategy of combining activity of
two defensin genes into a single-fusion gene may
have promising utility for biotechnological
applications.
Comments
Radish plants are propagated primarily from seeds.
Selected References
Abdou IA, Abou-Zeid AA, El-Sherbeeny MR, Abou-El-
Gheat ZH (1972) Antimicrobial activities of Allium
sativum, Allium cepa, Raphanus sativus, Capsicum
frutescens, Eruca sativa, Allium kurrat on bacteria.
Qual Plant Mater Veg 22(1):29–35
Aerts AM, François IE, Meert EM, Li QT, Cammue BP,
Thevissen K (2007) The antifungal activity of
RsAFP2, a plant defensin from Raphanus sativus,
involves the induction of reactive oxygen species in
Candida albicans. J Mol Microbiol Biotechnol
13(4):243–247
Aerts AM, Carmona-Gutierrez D, Lefevre S, Govaert G,
François IE, Madeo F, Santos R, Cammue BP,
Thevissen K (2009) The antifungal plant defensin
RsAFP2 from radish induces apoptosis in a metacas-
pase independent way in Candida albicans. FEBS Lett
583(15):2513–2516
Afsharypuor S, Balam MH (2005) Volatile constituents of
Raphanus sativus L. var. niger seeds. Essent Oil Res
17(4):440–441
Ahmad F, Hasan I, Chishti DK, Ahmad H (2013)
Antibacterial activity of Raphanus sativus Linn. seed
extract. Glob J Med Res 12(11):25–28
Ahn MJ, Koh RK, Kim GO, Shin TK (2013) Aqueous
extract of purple Bordeaux radish, Raphanus sativus
861
L. ameliorates ethanol-induced gastric injury in rats.
Orient Pharm Exp Med 13(4):247–252
Algasoumi S, Al-Yahya M, Al-Howiriny T, Rafatullah S
(2008) Gastroprotective effect of radish “Raphanus sativus
L”. on experimental models. Farmacia 56(2):204–214
Al-Shebaz IA (1985) The genera of Brassicaceae in the
southeastern United States. J Arnold Arboretum
66:279–351
Alves AL, De Samblanx GW, Terras FR, Cammue BP,
Broekaert WF (1994) Expression of functional
Raphanus sativus antifungal protein in yeast. FEBS
Lett 348(3):228–232
Anwar R, Mubasher Ahmad M (2006) Studies of
Raphanus sativus as hepato protective agent. J Med
Sci 6(4):662–665
Arai Y, Watanabe S, Kimira M, Shimoi K, Mochizuki R,
Kinae N (2000) Dietary intakes of flavonols, flavones
and isoflavones by Japanese women and the inverse
correlation between quercetin intake and plasma LDL
cholesterol concentration. J Nutr 130(9):2243–2250
Aspart L, Cooke R, Delseny M (1979) Stability of polyad-
enylic and polyadenylated ribonucleic acids in radish
(Raphanus sativus) seedlings. Biochim Biophys Acta
564(1):43–54
Aspart-Pascot L, Delseny M, Guitton Y (1976) Occurrence
of nucleoside polyphosphates in rapidly labelled RNA
preparations from radish seedlings (Raphanus sati-
vus.). Planta 131(3):275–278
Bach TJ, Rogers DH, Rudney H (1986) Detergent-
solubilization, purification, and characterization of
membrane-bound 3-hydroxy-3-methylglutaryl-coen-
zyme A reductase from radish seedlings. Eur J
Biochem 154(1):103–111
Baek SH, Park M, Suh JH, Choi HS (2008) Protective
effects of an extract of young radish (Raphanus sati-
vus L) cultivated with sulphur (sulfur-radish extract)
and of sulforaphane on carbon tetrachloride induced
hepatotoxicity. Biosci Biotechnol Biochem
72(5):1176–1182
Barillari J, Cervellati R, Paolini M, Tatibouët A, Rollin P,
Iori R (2005) Isolation of 4-methylthio-3-butenyl glu-
cosinolate from Raphanus sativus sprouts (kaiware
daikon) and its redox properties. J Agric Food Chem
53(26):9890–9896
Barillari J, Cervellati R, Costa S, Guerra MC, Speroni E,
Utan A, Iori R (2006) Antioxidant and choleretic prop-
erties of Raphanus sativus L. sprout (Kaiware Daikon)
extract. J Agric Food Chem 54(26):9773–9778
Barillari J, Iori R, Papi A, Orlandi M, Bartolini G,
Gabbanini S, Pedulli GF, Valgimigli L (2008) Kaiware
Daikon (Raphanus sativus L.) extract: a naturally mul-
tipotent chemopreventive agent. J Agric Food Chem
56(17):7823–7830
Beevi SS, Mangamoori LN, Dhand V, Ramakrishna DS
(2009) Isothiocyanate profile and selective antibacte-
rial activity of root, stem, and leaf extracts derived
from Raphanus sativus L. Foodborne Pathog Dis
6(1):129–136
Beevi SS, Mangamoori LN, Subathra M, Edula JR
(2010a) Hexane extract of Raphanus sativus L. roots
862
inhibits cell proliferation and induces apoptosis in
human cancer cells by modulating genes related to
apoptotic pathway. Plant Foods Hum Nutr
65(3):200–209
Beevi SS, Narasu ML, Gowda BB (2010b) Polyphenolics
profile, antioxidant and radical scavenging activity of
leaves and stem of Raphanus sativus L. Plant Foods
Hum Nutr 65(1):8–17
Beevi SS, Mangamoori LN, Gowda BB (2012)
Polyphenolics profile and antioxidant properties of
Raphanus sativus L. Nat Prod Res 26(6):557–563
Berger S, Menudier A, Julien R, Karamanos Y (1995)
Endo-N-acetyl-beta-D-glucosaminidase and peptide-
N4-(N-acetyl-glucosaminyl) asparagine amidase
activities during germination of Raphanus sativus.
Phytochemistry 39(3):481–487
Bilyk A, Sapers GM (1985) Distribution of quercetin and
kaempferol in lettuce, kale, chive, garlic chive, leek,
horseradish, red radish, and red cabbage tissues. J
Agric Food Chem 33(2):226–228
Blažević I, Mastelić J (2009) Glucosinolate degradation
products and other bound and free volatiles in the
leaves and roots of radish (Raphanus sativus L.). Food
Chem 113(1):96–102
Bodson MJ, Outlaw WH, Silvers SH (1991) Malate con-
tent of picoliter samples of Raphanus sativus cyto-
plasm. J Histochem Cytochem 39(4):435–440
Brandl W, Herrmann K, Grothjahn L (1984)
Hydroxycinnamoyl esters of malic acid in small radish
Raphanus sativus L. var. sativus. Z Naturforsch Teil C
39:515–520
Cai QY, Mo CH, Wu QT, Zeng QY (2008) Polycyclic aro-
matic hydrocarbons and phthalic acid esters in the
soil-radish (Raphanus sativus) system with sewage
sludge and compost application. Bioresour Technol
99(6):1830–1836
Capecka E (1998) Changes in thiocyanate content in some
radish Raphanus sativus L. cultivars during hypocotyl-
root growth. Acta Physiol Plant 20(2):135–142
Carlson DG, Daxenbiehler ME, VanEtten CH, Hill CB,
Williams PH (1985) Glucosinolates in radish cultivars.
J Amer Soc Hort Sci 110(5):634–638
Castro-Torres IG, Naranjo-Rodríguez EB, Domínguez-
Ortíz MÁ, Gallegos-Estudillo J, Saavedra-Vélez MV
(2012) Antilithiasic and hypolipidaemic effects of
Raphanus sativus L. var. niger on mice fed with a
lithogenic diet. J Biomed Biotechnol 2012:161205
Castro-Torres IG, De la O-Arciniega M, Gallegos-
Estudillo J, Naranjo-Rodríguez EB, Domínguez-Ortíz
MÁ (2014) Raphanus sativus L. var niger as a source
of phytochemicals for the prevention of cholesterol
gallstones. Phytother Res 28(2):167–171
Chandra AK, Mukhopadhyay S, Lahari D, Tripathy S
(2004) Goitrogenic content of Indian cyanogenic plant
foods & their in vitro anti-thyroidal activity. Indian J
Med Res 119(5):180–185
Chandra AK, Mukhopadhyay S, Ghosh D, Tripathy S
(2006) Effect of radish (Raphanus sativus Linn.) on
thyroid status under conditions of varying iodine
intake in rats. Indian J Exp Biol 44(8):653–661
Brassicaceae
Chaturvedi P (2008) Inhibitory response of Raphanus
sativus on lipid peroxidation in albino rats. Evid Based
Complement Alternat Med 5(1):55–59
Chaturvedi P, Machacha CN (2007) Efficacy of Raphanus
sativus in the treatment of paracetamol-induced hepa-
totoxicity in albino rats. Br J Biomed Sci
64(3):105–108
Chaturvedi P, George S, Machacha CN (2007) Protective
role of Raphanus sativus root extract on paracetamol-
induced hepatotoxicity in albino rats. Int J Vitam Nutr
Res 77(1):41–45
Chen SH, Lv GY, Zhang XD, Liu XY, Zhang H, Zhu YW,
Wu Y, Liu SY, Ni ZN (2007) Anti-hypertensive effects
of laiju extract in two different rat models. Asia Pac J
Clin Nutr 16(Suppl 1):309–312
Chinese Pharmacopoeia Commission (2010)
Pharmacopoeia of the People’s Republic of China, vol
1. Chinese Medical Science Press, Beijing, p 255
Chong C, Bible B (1975) Influence of seed on thiocyanate
content of radishes. J Sci Food Agric 26(1):105–108
Cocucci M, Negrini N (1988) Changes in the levels of
calmodulin and of a calmodulin inhibitor in the early
phases of radish (Raphanus sativus L.) seed germina-
tion: effects of aba and fusicoccin. Plant Physiol
88(3):910–914
Dahlbender B, Strack D (1984) Nitrogen nutrition and
the accumulation of free and sinapoyl-bound malic
acid in Raphanus sativus cotyledons. Planta
161(2):142–147
Damiani E, Aloia AM, Priore MG, Nardulli S, Ferrannini
A (2011) Generalized urticaria after ingestion of
Raphanus sativus. Ann Allergy Asthma Immunol
106(2):168
Dande P, Vaidya A, Arora P (2014) Laxative activity of
Raphanus sativus L. leaf. Asian J Pharm Clin Res
7(2):120–124
Devaraj VC, Krishna BG, Viswanatha GL (2011)
Simultaneous determination of quercetin, rutin and
kaempferol in the leaf extracts of Moringa oleifera
Lam. and Raphanus sativus Linn. by liquid
chromatography-tandem mass spectrometry. Zhong
Xi Yi Jie He Xue Bao 9(9):1022–1030
Domingos AK, Saad EB, Wilhelm HM, Ramos LP (2008)
Optimization of the ethanolysis of Raphanus sativus
(L. Var.) crude oil applying the response surface meth-
odology. Bioresour Technol 99(6):1837–1845
Ediage EN, Di Mavungu JD, Scippo ML, Schneider YJ,
Larondelle Y, Callebaut A, Robbens J, Van Peteghem
C, De Saeger S (2011) Screening, identification and
quantification of glucosinolates in black radish
(Raphanus sativus L. niger) based dietary supple-
ments using liquid chromatography coupled with a
photodiode array and liquid chromatography-mass
spectrometry. J Chromatogr A 1218(28):4395–4405
Elivra B, Tatiana B (2012) Investigations of thioglyco-
sides Raphanus sativus. Adv Res Sci Areas
3(7):1424–1425
Entsch B, Letham DS (1979) Enzymic glucosylation of
the cytokinin 6-aminobenzylaminopurine. Plant Sci
Lett 14(2):205–212
Raphanus raphanistrum subsp. sativus
Fant F, Vranken W, Broekaert W, Borremans F (1998)
Determination of the three-dimensional solution struc-
ture of Raphanus sativus antifungal protein 1 by 1H
NMR. J Mol Biol 279(1):257–270
Faye L, Berjonneau C (1979) Evidence for the glycopro-
tein nature of radish beta-fructosidase. Biochimie
61(1):51–59
Faye L, Mouatassim B, Ghorbel A (1986) Cell wall and
cytoplasmic isozymes of radish beta-fructosidase have
different n-linked oligosaccharides. Plant Physiol
80(1):27–33
Gao RC, Jing P, Ruan SY, Zhang YF, Zhao SJ, Cai Z, Qian
BJ (2014) Removal of off-flavours from radish
(Raphanus sativus L.) anthocyanin-rich pigments
using chitosan and its mechanism(s). Food Chem
146:423–428
Ghayur MN, Gilani AH (2005) Gastrointestinal stimula-
tory and uterotonic activities of dietary radish leaves
extract are mediated through multiple pathways.
Phytother Res 19(9):750–755
Ghayur MN, Gilani AH (2006) Radish seed extract
mediates its cardiovascular inhibitory effects via mus-
carinic receptor activation. Fundam Clin Pharmacol
20(1):57–63
Ghayur MN, Gilani AH (2012) Contractile effect of radish
and betel nut extracts on rabbit gallbladder. J
Complement Integr Med 9, 3
Ghayur MN, Gilani AH, Houghton PJ (2005) Species dif-
ferences in the gut stimulatory effects of radish seeds.
J Pharm Pharmacol 57(11):1493–1501
Gilani AH, Ghayur MN (2004) Pharmacological basis for
the gut stimulatory activity of Raphanus sativus
leaves. J Ethnopharmacol 95(2–3):169–172
Gräwe W, Bachhuber P, Mock HP, Strack D (1992)
Purification and characterization of
sinapoylglucose:malate sinapoyltransferase from
Raphanus sativus L. Planta 187(2):236–241
Greer MA (1950) Nutrition and goiter. Physiol Rev
30:513–548
Greuter Q, Burdet (2001) Raphanus Linnaeus. In: Wu ZY,
Raven PH (eds) Flora of China, vol 8, Brassicaceae
through Saxifragaceae. Science Press/Missouri
Botanical Garden Press, Beijing/St. Louis
Guisti MM, Ghanadan H, Wrolstad RE (1998) Elucidation
of the structure and conformation of red radish
(Raphanus sativus) anthocyanins using one- and two-
dimensional nuclear magnetic resonance techniques. J
Agric Food Chem 46(12):4858–4863
Gutiérrez RMP, Perez RL (2004) Raphanus sativus
(radish): their chemistry and biology. Sci World J
4:811–837
Hanlon PR, Barnes DM (2011) Phytochemical composi-
tion and biological activity of 8 varieties of radish
(Raphanus sativus L.) sprouts and mature taproots. J
Food Sci 76(1):C185–C192
Hanlon PR, Webber DM, Barnes DM (2007) Aqueous
extract from Spanish black radish (Raphanus sativus
L. var. niger) induces detoxification enzymes in the
HepG2 human hepatoma cell line. J Agric Food Chem
55(16):6439–6446
863
Hanlon PR, Robbins MG, Hammon LD, Barnes DM
(2009) Aqueous extract from the vegetative portion of
Spanish black radish (Raphanus sativus L. var. niger)
induces detoxification enzyme expression in HepG2
cells. J Funct Foods 1:356–365
Hara M, Fujii Y, Sasada Y, Kuboi T (2000) cDNA cloning
of radish (Raphanus sativus) myrosinase and tissue-
specific expression in root. Plant Cell Physiol
41(10):1102–1109
Hara M, Ito F, Asai T, Kuboi T (2009) Variation in amy-
lase activities in radish (Raphanus sativus) cultivars.
Plant Foods Hum Nutr 64(3):188–192
Hara M, Torazawa D, Asai T, Takahashi I (2011) Variations
in the soluble sugar and organic acid contents in radish
(Raphanus sativus L.) cultivars. Int J Food Sci Technol
46(11):2387–2392
Harnly JM, Doherty R, Beecher GR, Holden JM,
Haytowitz DB, Bhagwat S, Gebhardt S (2006)
Flavonoid content of U.S. fruits, vegetables, and nuts.
J Agric Food Chem 54(26):9966–9977
Hase T, Hasegawa K (1982) Raphanusol A, a new growth
inhibitor from Sakurajima radish seedlings.
Phytochemistry 21(5):1021–1022
Hase T, Koreeda M, Hasegawa K (1983) A growth inhibi-
tor, 2-thioxothiazolidine-4-carboxylic acid from
Sakurajima radish seedlings. Phytochemistry 22(5):
1275–1276
Hasegawa K, Hase T (1981) Raphanusol B: a growth
inhibitor of light-grown radish seedlings. Plant Cell
Physiol 22(2):303–306
Hasegawa K, Miyamoto K (1978) Light growth inhibition
and growth inhibitors in Sakurajima radish seedlings.
Plant Cell Physiol 19(6):1077–1083
Hasegawa K, Miyamoto K (1980) Raphanusol A: a new
growth inhibitor of light-grown radish seedlings. Plant
Cell Physiol 21(2):363–366
Hasegawa K, Shiihara S, Iwagawa T, Hase T (1982)
Isolation and identification of a new growth inhibitor,
raphanusanin, from radish seedlings and its role in
light inhibition of hypocotyl growth. Plant Physiol
70(2):626–628
Hasegawa K, Noguchi H, Iwagawa T, Hase T (1986)
Phototropism in hypocotyls of radish i. isolation and
identification of growth inhibitors, cis-and trans-
raphanusanins and raphanusamide, involved in photot-
ropism of radish hypocotyls. Plant Physiol 81(4):
976–979
Hata K, Tanaka M, Tsumuraya Y, Hashimoto Y (1992)
α-L-Arabinofuranosidase from radish (Raphanus sati-
vus L.) seeds. Plant Physiol 100(1):388–396
Herrmann K (1989) Occurrence and content of hydroxy-
cinnamic and hydroxybenzoic acid compounds in
foods. Crit Rev Food Sci Nutr 28:315–347
Hertog MGL, Hollman PCH, Katan MB (1992) Content
of potentially anticarcinogenic flavonoids of 28 vege-
tables and fruits commonly consumed in The
Netherlands. J Agric Food Chem 40(12):2379–2383
Huxley A, Griffiths M, Levy M (eds) (1997) The new
royal horticultural society dictionary of gardening,
volume 2 (D–K). The Stockton Press, New York
864
Ishii G, Saijo R (1987) Effect of season, soil type, sul-
phate level, mulching and plant density on isothiocya-
nate content in radish root juice. J Jpn Soc Hort Sci
56(3):313–320
Ishii G, Saijo R, Mizutani J (1989a) A quantitative deter-
mination of 4-methylthio-3-butenyl glucosinolate in
daikon (Raphanus sativus L.) roots by gas liquid chro-
matography. J Jpn Soc Hort Sci 58(2):339–344
Ishii G, Saijo R, Nagata M (1989b) The difference of glu-
cosinolate content in different cultivar of daikon roots
(Raphanus sativus L.). Nippon Shokuhin Kogyo
Gakkaishi 36(9):739–742
Ishijima J, Nagasaki N, Maeshima M, Miyano M (2007)
RVCaB, a calcium-binding protein in radish vacuoles,
is predominantly an unstructured protein with a poly-
proline type II helix. J Biochem 142(2):201–211
Ivanovics G, Horvath S (1947) Raphanin, an antibacterial
principle of the radish (Raphanus sativus). Nature
160(4061):297–298
Jakmatakul R, Suttisri R, Tengamnuay P (2009)
Evaluation of antityrosinase and antioxidant activities
of Raphanus sativus root: comparison between freeze-
dried juice and methanolic extract. Thai J Pharm Sci
33(1):22–30
Janjua S, Shahid M, Fakhir-i-Abbas (2013) Phytochemical
analysis and in vitro antibacterial activity of root peel
extract of Raphanus sativus. L. var niger. Adv Med
Plant Res 1(1):1–7
Jing P, Zhao SJ, Ruan SY, Xie ZH, Dong Y, Yu LI (2012)
Anthocyanin and glucosinolate occurrences in the
roots of Chinese red radish (Raphanus sativus L.), and
their stability to heat and pH. Food Chem
133(4):1569–1576
Jwanny EW, El-Sayed ST, Rashad MM, Mahmoud AE,
Abdallah NM (1995) Myrosinase from roots of
Raphanus sativus. Phytochemistry 39(6):1301–1303
Kamble S, Ahmed Z, Ramabhimaiaha S, Patil P (2013)
Anti-inflammatory activity of Raphanus sativus L in
acute and chronic experimental models in albino rats.
Biomed Pharmacol J 6(2):315–320
Karri V, Bharadwaja KP (2013) Tandem combination of
Trigonella foenum-graecum defensin (Tfgd2) and
Raphanus sativus antifungal protein (RsAFP2) gener-
ates a more potent antifungal protein. Funct Integr
Genomics 13(4):435–443
Katsuzaki H, Miyahara Y, Ota M, Imai K, Komiya T
(2004) Chemistry and antioxidative activity of hot
water extract of Japanese radish (daikon). Biofactors
21(1–4):211–214
Khanal P, Karmacharya A, Sharma S, Nepal AK, Shrestha K
(2014) Biotechnological production of inducible
defense-related proteins in edible radish (Raphanus sati-
vus) found in Nepal. J Clin Diagn Res 8(1):112–115
Kikuchi Y, Saika H, Yuasa K, Nagahama M, Tsuji A
(2008) Isolation and biochemical characterization of
two forms of RD21 from cotyledons of daikon radish
(Raphanus sativus). J Biochem 144(6):789–798
Kim SS, Lee DJ (2005) Purification and characterization
of a cationic peroxidase Cs in Raphanus sativus. J
Plant Physiol 162(6):609–617
Brassicaceae
Kim HJ, Chen F, Wang X, Choi JH (2006) Effect of
methyl jasmonate on phenolics, isothiocyanate, and
metabolic enzymes in radish sprout (Raphanus sativus
L.). J Agric Food Chem 54(19):7263–7269
Kim WK, Kim JH, Jeong DH, Chun YH, Kim SH, Cho KJ,
Chang MJ (2011) Radish (Raphanus sativus L. leaf)
ethanol extract inhibits protein and mRNA expression
of ErbB(2) and ErbB(3) in MDA-MB-231 human
breast cancer cells. Nutr Res Pract 5(4):288–293
Kim KH, Moon EJ, Kim SY, Choi SU, Lee JH, Lee KR
(2014) 4-Methylthio-butanyl derivatives from the
seeds of Raphanus sativus and their biological evalua-
tion on anti-inflammatory and antitumor activities. J
Ethnopharmacol 151(1):503–508
Kjaer A, Madsen JO, Maeda Y, Ozawa Y, Uda Y (1978)
Volatiles in distillates of fresh radish of Japanese and
Kenyan origin. Agric Biol Chem 42(9):1715–1721
Kotake T, Tsuchiya K, Aohara T, Konishi T, Kaneko S,
Igarashi K, Samejima M, Tsumuraya Y (2006) An
alpha-L-arabinofuranosidase/beta-D-xylosidase from
immature seeds of radish (Raphanus sativus L.). J Exp
Bot 57(10):2353–2362
Lai XF, Cai FG, Wang WJ, Ma LQ, Zhong YX, Xu FC
(2006) Purification and characterization of two chitin-
binding proteins with lysozyme activity from roots of
Raphanus sativus. Zhi Wu Sheng Li Yu Fen Zi Sheng
Wu Xue Xue Bao 32(4):445–450 (In Chinese)
Laroche M, Aspart L, Delseny M, Penon P (1984)
Characterization of radish (Raphanus sativus) storage
proteins. Plant Physiol 74(3):487–493
Lee MY, Kim SS (1994) Characteristics of six isoperoxi-
dases from Korean radish root. Phytochemistry
35(2):287–290
Lee SO, Lee IS (2006) Induction of quinone reductase,
the phase 2 anticarcinogenic marker enzyme, in
Hepa1c1c7 cells by radish sprouts, Raphanus sativus
L. J Food Sci 71(2):S144–S148
Lee SW, Yang KM, Kim JK, Nam BH, Lee CM, Jeong
MH, Seo SY, Kim GY, Jo WS (2012) Effects of white
radish (Raphanus sativus) enzyme extract on hepato-
toxicity. Toxicol Res 28(3):165–172
Li YB, Jing S (2010) Determination of the total alkaloids
in Semen Raphani by acid dye colorimetry. Inf Trad
Chin Med 27:8–10 (In Chinese)
Li TY, Li TG, Zhang GX, Guo HY, Liu J, Li BG, Piao ZY,
Gai GG (2007) Experimental study of hypotensive
effect of soluble Semen Raphani alkaloids in sponta-
neous hypertensive rats. World J Integr Trad West Med
2(1):25–28 (In Chinese)
Li QY, Dai SJ, Chen SX, Yan XF (2008) Analysis of glu-
cosinolate composition and content in radish. Acta
Hort Sin 35(8):1205–1208 (In Chinese)
Lichtenthaler HK, Becker K (1975) The influence of con-
tinuous far-red and white light on prenyl chain synthe-
sis in plastids of Raphanus seedlings. Planta
122(3):255–258
Lin LZ, Sun J, Chen P, Harnly JA (2011) LC-PDA-ESI/
MSn identification of new anthocyanins in purple
Bordeaux radish (Raphanus sativus L. variety). J
Agric Food Chem 59(12):6616–6627
Raphanus raphanistrum subsp. sativus
Linscheid M, Wendisch D, Strack D (1980) The structure of
sinapic acid esters and their metabolism in cotyledons
of Raphanus sativus. Z Naturforsch C 35:907–914
Liu LF, Wang YX, Zhang XY, Liu XF (2002)
Determination of sinapine in Semen Raphani. Chin
Trad Patent Med 24:52–54 (In Chinese)
Liu YH, Murakami N, Wang LS, Zhang S (2008)
Preparative high-performance liquid chromatography
for the purification of natural acylated anthocyanins
from red radish (Raphanus sativus L.). J Chromatogr
Sci 46(8):743–746
Liu J, Dong N, Wang Q, Li J, Qian G, Fan H, Zhao G
(2014) Thermal degradation kinetics of anthocyanins
from Chinese red radish (Raphanus sativus L.) in
various juice beverages. Eur Food Res Technol
238(2):177–184
Lugasi A, Hovari J (2000) Flavonoid aglycons in foods of
plant origin I. Veg Acta Aliment 29:345–352
Lugasi A, Blázovics A, Hagymási K, Kocsis I, Kéry A
(2005) Antioxidant effect of squeezed juice from
black radish (Raphanus sativus L. var niger) in ali-
mentary hyperlipidaemia in rats. Phytother Res
19(7):587–591
Mabberley DJ (2008) Mabberley’s plant-book: a portable
dictionary of plants, their classification and uses, 3rd
edn. Cambridge University Press, Cambridge
Makhkamov GM, Latipov A (1965) Effect of natural goi-
trogenic compounds on hormone formation in thyroid
glands of rats. Uzbeksk Biol Zh 9:10–13
Marquardt P, Classen HG, Schumacher KA (1976)
N-Methylphenethylamine, an indirect sympathicomi-
metic agent in vegetables. Arzneimittelforschung
26(11):2001–2003 (In German)
Martínez-Villaluenga C, Frías J, Gulewicz P, Gulewicz K,
Vidal-Valverde C (2008) Food safety evaluation of
broccoli and radish sprouts. Food Chem Toxicol
46(5):1635–1644
Matsufuji H, Kido H, Misawa H, Yaguchi J, Otsuki T,
Chino M, Takeda M, Yamagata K (2007) Stability to
light, heat, and hydrogen peroxide at different pH val-
ues and DPPH radical scavenging activity of acylated
anthocyanins from red radish extract. J Agric Food
Chem 55(9):3692–3701
Matsuoka H, Takahashi A, Yanagi K, Uda Y (1997)
Antimicrobial action of 2-thioxo-3-
pyrrolidinecarbaldehyde, a major thiolactam com-
pound generated from the pungent principle of radish
in an aqueous medium. Food Sci Technol Int (Tokyo)
3(4):353–356
Mayer C (1981) Vegetarian medicines. Mayerbooks,
Glenwood, p. 20
Mezencev R, Kutschy P, Salayova A, Updegrove T,
McDonald JF (2009) The design, synthesis and anti-
cancer activity of new nitrogen mustard derivatives of
natural indole phytoalexin 1-methoxyspirobrassinol.
Neoplasma 56(4):321–330
Misawa H, Tsumuraya Y, Kaneko Y, Hashimoto Y (1996)
[alpha]-L-fucosyltransferases from radish primary
roots. Plant Physiol 110(2):665–673
865
Mitchell JC, Jordan WP (1974) Allergic contact dermatitis
from the radish, Raphanus sativus. Br J Dermatol
91(2):183–189
Mock HP, Strack D (1993) Energetics of the uridine
5′-diphosphoglucose: hydroxycinnamic acid acyl-
glucosyltransferase reaction. Phytochemistry
32(3):575–579
Mohammed NHSH, Abelgasim AI, Mohammed AH
(2008) Protective effect of Raphanus sativus against
carbon tetrachloride induced hepatotoxicity in wistar
albino rats. J Pharmacol Toxicol 3(4):272–278
Montaut S, Barillari J, Iori R, Rollin P (2010)
Glucoraphasatin: chemistry, occurrence, and biologi-
cal properties. Phytochemistry 71(1):6–12
Moreland DE, Hussey GG, Shriner CR, Farmer FS (1974)
Adenosine phosphates in germinating radish (Raphanus
sativus L.) seeds. Plant Physiol 54(4):560–563
Muminova BA, Batirov EK, Yuldashev MP, Inamova ZG
(2006) Kaempferol glycosides from Allium cepa and
Raphanus sativus. Chem Nat Comp 42(1):110–111
Naghibi F, Pourmorad F, Honary S, Shamsi M (2003)
Decontamination of water polluted with phenol using
Raphanus sativus root. Iranian J Pharm Res
2(1):29–32
Nakamura Y, Iwahashi T, Tanaka A, Koutani J, Matsuo T,
Okamoto S, Sato K, Ohtsuki K (2001) 4-(Methylthio)-
3-butenyl isothiocyanate, a principal antimutagen in
daikon (Raphanus sativus; Japanese white radish). J
Agric Food Chem 49(12):5755–5760
Nakamura Y, Nakamura K, Asai Y, Wada T, Tanaka K,
Matsuo T, Okamoto S, Meijer J, Kitamura Y,
Nishikawa A, Park EY, Sato K, Ohtsuki K (2008)
Comparison of the glucosinolate-myrosinase systems
among daikon (Raphanus sativus, Japanese white rad-
ish) varieties. J Agric Food Chem 56(8):2702–2707
Nakano Y, Okawa S, Yamauchi T, Koizumi Y, Sekiya J
(2006) Purification and properties of soluble and
bound gamma-glutamyltransferases from radish coty-
ledon. Biosci Biotechnol Biochem 70(2):369–376
Nakayama M, Yamane H, Yokota T, Yamaguchi I,
Murofushi N, Takahashi N, Nishijima T, Katsura N,
Nonaka M, Gaskin P, MacMillan J, Mander LN, Chu
A (1990) Endogenous gibberellins in mature seed of
Raphanus sativus L. cv. Taibyo-sobutori. Agric Biol
Chem 54(3):837–840
Nakayama M, Yamane H, Nojiri H, Yokota T, Yamaguchi I,
Murofushi N, Takahashi N, Nishijima T, Koshioka M,
Katsura N, Nonaka M (1995) Qualitative and quantita-
tive analysis of endogenous gibberellins in Raphanus
sativus L. during cold treatment and the subsequent
growth. Biosci Biotechnol Biochem 59(6):1121–1125
Nielsen JK, Olsen O, Pedersen LH, Sørensen H (1984)
2-O-(p-Coumaroyl)-l-malate, 2-O-caffeoyl-l-malate
and 2-O-feruloyl-l-malate in Raphanus sativus.
Phytochemistry 23(8):1741–1743
Nishijima T, Koshioka M, Yamazaki H, Miura H, Mander
LN (1995) Endogenous gibberellins and bolting in
cultivars of Japanese radish. Acta Hort (ISHS)
394:199–206
866
N’jai AU, Kemp MQ, Metzger BT, Hanlon PR, Robbins
M, Czuyprynski C, Barnes DM (2012) Spanish black
radish (Raphanus sativus L. var. niger) diet enhances
clearance of DMBA and diminishes toxic effects on
bone marrow progenitor cells. Nutr Cancer
64(7):1038–1048
Nunes DL, Franca AS, Oliveira LS (2011) Use of
Raphanus sativus L. press cake, a solid residue from
biodiesel processing, in the production of adsorbents
by microwave activation. Environ Technol
32(9–10):1073–1083
Nurmann G, Strack D (1979) Sinapine esterase. 1.
Characterization of sinapine esterase from cotyledons
of Raphanus sativus. Z Naturforsch C 34:715–720
Obata S, Nishimura M, Nagai K, Sakihama N, Shin M
(1995) Four ferredoxins from Japanese radish leaves.
Arch Biochem Biophys 316(2):797–802
Ochse JJ, Bakhuizen van den Brink RC (1980) Vegetables
of the Dutch Indies, 3rd edn. Ascher & Co.,
Amsterdam, 1016 pp
Okano K, Asano J, Ishii G (1990) Contents of pungent
principle in roots of Japanese radish (Raphanus sati-
vus L.) cultivars. J Jpn Soc Hort Sci 59(3):551–558
Orlovskaya T, Sedin A, Malikova M (2013) Carbohydrates
from seeds of Raphanus sativus and Brassica napus.
Chem Nat Comp 49(2):327–328
Otsuki T, Matsufuji H, Takeda M, Toyoda M, Goda Y
(2002) Acylated anthocyanins from red radish
(Raphanus sativus L.). Phytochemistry 60(1):79–87
Papi A, Orlandi M, Bartolini G, Barillari J, Iori R, Paolini
M, Ferroni F, Grazia Fumo M, Pedulli GF, Valgimigli
L (2008) Cytotoxic and antioxidant activity of
4-methylthio-3-butenyl isothiocyanate from Raphanus
sativus L. (Kaiware Daikon) sprouts. J Agric Food
Chem 56(3):875–883
Park NI, Xu H, Li X, Jang IH, Park S, Ahn GH, Lim YP,
Kim SJ, Park SU (2011) Anthocyanin accumulation
and expression of anthocyanin biosynthetic genes in
radish (Raphanus sativus). J Agric Food Chem
59(11):6034–6039
Pedrero Z, Madrid Y, Cámara C (2006) Selenium species
bioaccessibility in enriched radish (Raphanus sati-
vus): a potential dietary source of selenium. J Agric
Food Chem 54(6):2412–2417
Phelan JR, Allen A, Vaughan JG (1984) Myrosinase in
Raphanus sativus L. J Exp Bot 35(10):1558–1564
Piluek K, Beltran MM (1994) Raphanus sativus L. In:
Siemonsma JS, Piluek K (eds) Plant resources of
south-east Asia, vol 8, Vegetables. Prosea Foundation,
Bogor, pp 233–237
Prahoveanu E, Eşanu V (1987) Immunomodulation with
natural products. I. Effect of an aqueous extract of
Raphanus sativus niger on experimental influenza
infection in mice. Virologie 38(2):115–120 (In French)
Prahoveanu E, Eşanu V (1990) The effects of aqueous
extracts of Raphanus niger on an experimental influ-
enza infection in mice and on the enzyme polymor-
phism in lung tissue extracts. Rev Roum Virol
41(2):113–117 (In French)
Brassicaceae
Rakhimov MM, Akhmedzhanov P, Babaev MU, Kkhole
B, Mad’iarov SP (1981) Properties of phospholipase
D from Raphanus sativus. Biokhimiia 46(2):240–249
(In Russian)
Roh SS, Park SB, Park SM, Choi BW, Lee MH, Hwang
YL, Kim CH, Jeong HA, Kim CD, Lee JH (2013) A
novel compound rasatiol isolated from Raphanus sati-
vus has a potential to enhance extracellular matrix syn-
thesis in dermal fibroblasts. Ann Dermatol
25(3):315–320
Sakakibara H, Honda Y, Nakagawa S, Ashida H,
Kanazawa K (2003) Simultaneous determination of all
polyphenols in vegetables, fruits, and teas. J Agric
Food Chem 51(3):571–581
Sakoda M, Hase T, Hasegawa K (1990) A growth inhibi-
tor, 3-(E)-(methylthio) methylene-2-pyrrolidinethione
from light-grown radish seedlings. Phytochemistry
29(4):1031–1032
Sakoda M, Hasegawa K, Ishizuka K (1991) The occur-
rence in plants of the growth inhibitors, the raphanusa-
nins. Phytochemistry 30(1):57–61
Salah-Abbès JB, Abbès S, Houas Z, Abdel-Wahhab MA,
Oueslati R (2008a) Zearalenone induces immunotox-
icity in mice: possible protective effects of radish
extract (Raphanus sativus). J Pharm Pharmacol
60(6):761–770
Salah-Abbès JB, Abbès S, Ouanes Z, Houas Z, Abdel-
Wahhab MA, Bacha H, Oueslati R (2008b) Tunisian
radish extract (Raphanus sativus) enhances the anti-
oxidant status and protects against oxidative stress
induced by zearalenone in Balb/c mice. J Appl Toxicol
28(1):6–14
Salah-Abbès JB, Abbès S, Abdel-Wahhab MA, Oueslati R
(2009a) Raphanus sativus extract protects against
Zearalenone induced reproductive toxicity, oxidative
stress and mutagenic alterations in male Balb/c mice.
Toxicon 53(5):525–533
Salah-Abbès JB, Abbès S, Haous Z, Oueslati R (2009b)
Raphanus sativus extract prevents and ameliorates
zearalenone-induced peroxidative hepatic damage in
Balb/c mice. J Pharm Pharmacol 61(11):1545–1554
Salah-Abbès JB, Abbès S, Ouanes Z, Abdel-Wahhab MA,
Bacha H, Oueslati R (2009c) Isothiocyanate from the
Tunisian radish (Raphanus sativus) prevents genotox-
icity of Zearalenone in vivo and in vitro. Mutat Res
677(1–2):59–65
Salah-Abbès JB, Abbès S, Abdel-Wahhab M, Oueslati R
(2010a) Immunotoxicity of zearalenone in Balb/c
mice in a high subchronic dosing study counteracted
by Raphanus sativus extract. Immunopharmacol
Immunotoxicol 32(4):628–636
Salah-Abbès JB, Abbès S, Abdel-Wahhab MA, Oueslati R
(2010b) In-vitro free radical scavenging, antiprolifera-
tive and anti-zearalenone cytotoxic effects of
4-(methylthio)-3-butenyl isothiocyanate from
Tunisian Raphanus sativus. J Pharm Pharmacol
62(2):231–239
Salah-Abbès JB, Abbès S, Zohra H, Oueslati R (2014)
Tunisian radish (Raphanus sativus) extract prevents
Raphanus raphanistrum subsp. sativus
cadmium-induced immunotoxic and biochemical
alterations in rats. J Immunotoxicol (in press)
Sang JP, Minchinton IR, Johnstone PK, Truscott RJW
(1984) Glucosinolate profiles in the seed, root and leaf
tissue of cabbage, mustard, rapeseed, radish and
Swede. Can J Plant Sci 64:77–93
San Juan MEC, Jumala RS, Niasca KHG (2012)
Phytochemical screening, isolation and structure elu-
cidation of the radish (Raphanus sativus Linn.) bulb
ethanolic extract using gas chromatography mass
spectrometry (GC-MS). Univ Immaculate Conception
Res J 18(1):237–247
Schmidtlein H, Herrmann K (1975) On phenolic acids of
vegetables. I. Hydroxycinnamic acids and hydroxy-
benzoic acids of Brassica-species and leaves of other
Cruciferae. Z Lebensm Unters Forsch 159(3):139–148
(In German)
Scholl C, Eshelman BD, Barnes DM, Hanlon PR (2011)
Raphasatin is a more potent inducer of the detoxifica-
tion enzymes than its degradation products. J Food Sci
76(3):C504–C511
Sekimata M, Ogura K, Tsumuraya Y, Hashimoto Y,
Yamamoto S (1989) A beta-galactosidase from radish
(Raphanus sativus L.) seeds. Plant Physiol
90(2):567–574
Sgherri C, Cosi E, Navari-Izzo F (2003) Phenols and anti-
oxidative status of Raphanus sativus grown in copper
excess. Physiol Plant 118(1):21–28
Sham TT, Yuen ACY, Ng YF, Chan CO, Mok DKW,
Chan SW (2013) A review of the phytochemistry and
pharmacological activities of Raphani Semen. Evid
Based Complement Alternat Med 2013: Article ID
636194
Sharma V, Strack D (1985) Vacuolar localization of
1-sinapolglucose: L-malate sinapoyltransferase in
protoplasts from cotyledons of Raphanus sativus.
Planta 163(4):563–568
Shikita M, Fahey JW, Golden TR, Holtzclaw WD, Talalay
P (1999) An unusual case of ‘uncompetitive activa-
tion’ by ascorbic acid: purification and kinetic proper-
ties of a myrosinase from Raphanus sativus seedlings.
Biochem J 341:725–732
Shukla S, Chatterji S, Mehta S, Rai PK, Singh RK, Yadav
DK, Watal G (2011a) Antidiabetic effect of Raphanus
sativus root juice. Pharm Biol 49(1):32–37
Shukla S, Chatterji S, Yadav DK, Watal G (2011b)
Antimicrobial efficacy of Raphanus sativus root juice.
Int J Pharm Pharm Sci 3:89–92
Sipos P, Hagymasi K, Lugasi A, Feher E, Blazovics A
(2002) Effects of black radish root (Raphanus sativus
L. var. niger) on the colon mucosa in rats fed a fat rich
diet. Phytother Res 16(7):677–679
Song K, Osborn TC, Williams PH (1990) Brassica taxon-
omy based on nuclear restriction fragment length
polymorphisms (RFLPs): 3. Genome relationships in
Brassica and related genera and the origin of B. olera-
cea and B. rapa (syn. campestris). Theor Appl Genet
79(4):497–506
867
Stöhr H, Herrmann K (1975) On the phenolic acids of
vegetables. III. Hydroxycinnamic acids and hydroxy-
benzoic acids of root vegetables. Z Lebensm Unters
Forsch 159(4):218–224 (In German)
Strack D (1977) Sinapic acid ester fluctuations in cotyle-
dons of Raphanus sativus. Z Pflanzenphysiol
84(2):139–145
Strack D (1981) Sinapine as a supply of choline for the
biosynthesis of phosphatidylcholine in Raphanus sati-
vus var sativus cultivar saxa seedlings. Z Naturforsch
C 4:215–221
Strack D (1982) Development of 1-O-sinapoyl-β-D-
glucose: l-malate sinapoyltransferase activity in coty-
ledons of red radish (Raphanus sativus L. var. sativus).
Planta 155:31–36
Strack D, Sharma V (1985) Vacuolar localization of the
enzymatic synthesis of hydroxycinnamic acid esters of
malic acid in protoplasts from Raphanus sativus
leaves. Physiol Plant 65:45–50
Strack D, Tkotz N, Klug M (1978) Phenylpropanoid
metabolism in cotyledons of Raphanus sativus and the
effect of competitive in vivo inhibition of
l-phenylalanine ammonia-lyase (PAL) by hydroxyl-
amine derivatives. Z Pflanzenphysiol 89:343–353
Strack D, Pieroth M, Scharf H, Sharma V (1985)
Tissue distribution of phenylpropanoid metabolism
in cotyledons of Raphanus sativus L. Planta
164(4):507–511
Strack D, Reinecke J, Takeuchi S (1986) Evidence for a
relationship between malate metabolism and activity
of 1-sinapoylglucose: L-malate sinapoyltransferase in
radish (Raphanus sativus L.) cotyledons. Planta
167(2):212–217
Streeter JG, Thompson JF (1972) Anaerobic accumula-
tion of gamma-aminobutyric acid and alanine in radish
leaves (Raphanus sativus, L.). Plant Physiol
49(4):572–578
Stuart GU (2013) Philippine alternative medicine. Manual
of some Philippine medicinal plants. http://www.stu-
artxchange.org/OtherHerbals.html
Suge H, Rappaport L (1968) Role of gibberellins in stem
elongation and flowering in radish. Plant Physiol
43(8):1208–1214
Suh SJ, Moon SK, Kim CH (2006) Raphanus sativus and
its isothiocyanates inhibit vascular smooth muscle
cells proliferation and induce G(1) cell cycle arrest. Int
Immunopharmacol 6(5):854–861
Takaya Y, Kondo Y, Furukawa T, Niwa M (2003)
Antioxidant constituents of radish sprout (Kaiware-
daikon), Raphanus sativus L. J Agric Food Chem
51(27):8061–8066
Tamura G, Iwasawa T, Masada M, Fukushima K (1976)
Some properties of cysteine synthase from radish
roots. Agric Biol Chem 40(3):637–638
Tamura S, Tsuji K, Yongzhen P, Ohnishi-Kameyama M,
Murakami N (2010) Six new acylated anthocyanins
from red radish (Raphanus sativus). Chem Pharm Bull
(Tokyo) 58(9):1259–1262
868
Tan P, Jiang HY, Lu WH (2005) A research review of
Raphani Semen. J Pract Trad Chin Med 21:254–255
Taniguchi H, Kobayashi-Hattori K, Tenmyo C, Kamei T,
Uda Y, Sugita-Konishi Y, Oishi Y, Takita T (2006)
Effect of Japanese radish (Raphanus sativus) sprout
(Kaiware-daikon) on carbohydrate and lipid metabo-
lisms in normal and streptozotocin-induced diabetic
rats. Phytother Res 20(4):274–278
Taniguchi H, Muroi R, Kobayashi-Hattori K, Uda Y, Oishi
Y, Takita T (2007) Differing effects of water-soluble
and fat-soluble extracts from Japanese radish (Raphanus
sativus) sprouts on carbohydrate and lipid metabolism
in normal and streptozotocin-induced diabetic rats. J
Nutr Sci Vitaminol (Tokyo) 53(3):261–266
Tatsuzawa F, Toki K, Saito N, Shinoda K, Shigihara A,
Honda T (2008) Anthocyanin occurrence in the root
peels, petioles and flowers of red radish (Raphanus
sativus L.). Dyes Pigments 79(1):83–88
Tatsuzawa F, Saito N, Toki K, Shinoda K, Shigihara A,
Honda T (2010) Acylated cyanidin 3-sophoroside-5-
glucosides from the purple roots of red radish
(Raphanus sativus L.)‘Benikanmi’. J Jpn Soc Hort Sci
79(1):103–107
Terras FR, Goderis IJ, Van Leuven F, Vanderleyden J,
Cammue BPA, Broekaert WF (1992a) In vitro antifun-
gal activity of a radish (Raphanus sativus L.) seed pro-
tein homologous to nonspecific lipid transfer proteins.
Plant Physiol 100(2):1055–1058
Terras FR, Schoofs HM, De Bolle MF, Van Leuven F,
Rees SB, Vanderleyden J, Cammue BP, Broekaert WF
(1992b) Analysis of two novel classes of plant antifun-
gal proteins from radish (Raphanus sativus L.) seeds.
J Biol Chem 267(22):15301–15309
Terras FR, Eggermont K, Kovaleva V, Raikhel NV,
Osborn RW, Kester A, Rees SB, Torrekens S, Van
Leuven F, Vanderleyden J, Cammue BPA, Broekaert
WF (1995) Small cysteine-rich antifungal proteins
from radish: their role in host defense. Plant Cell
7(5):573–588
The Plant List (2014) Raphanus raphanistrum subsp. sati-
vus. http://www.theplantlist.org/
Tkotz N, Strack D (1980) Enzymatic synthesis of
sinapoyl-L-malate from 1-sinapoylglucose and
L-malate by a protein preparation from Raphanus sati-
vus cotyledons. Z Naturforsch C 10:835–837
Tomè F, Campedelli L, Bellini E (1975) Distribution of
phenylalanine transaminase and phenylalanine
ammonia-lyase activities in etiolated and light irradi-
ated radish seedlings (Raphanus sativus L.).
Experientia 31(10):1119–1121
Tsuji A, Kikuchi Y, Ogawa K, Saika H, Yuasa K,
Nagahama M (2008) Purification and characterization
of cathepsin B-like cysteine protease from cotyledons
of daikon radish, Raphanus sativus. FEBS J
275(21):5429–5443
Tsumuraya Y, Hashimoto Y, Yamamoto S, Shibuya N
(1984) Structure of L-arabino-D-galactan-containing
glycoproteins from radish leaves. Carbohydr Res
134(2):215–228
Brassicaceae
Tsumuraya Y, Hashimoto Y, Yamamoto S (1987) An
L-arabino-D-galactan and an L-arabino-D-galactan-
containing proteoglycan from radish (Raphanus sati-
vus) seeds. Carbohydr Res 161(1):113–126
Tsumuraya Y, Ogura K, Hashimoto Y, Mukoyama H,
Yamamoto S (1988) Arabinogalactan-proteins from
primary and mature roots of radish (Raphanus sativus
L.). Plant Physiol 86(1):155–160
Uda Y, Matsuoka H, Kumagami H, Shima H, Maeda Y
(1993a) Stability and antimicrobial property of
4-methylthio-3-butenyl isothiocyanate, the pungent
principle in radish. Nippon Shokuhin Kogyo Gakkaishi
40(10):743–746
Uda Y, Matsuoka H, Shima H, Kumagami H, Maeda Y
(1993b) Antimicrobial activity of water-soluble prod-
ucts derived from radish mustard oil and identification
of an active component therein. Nippon Shokuhin
Kogyo Gakkaishi 40(11):801–806
U.S. Department of Agriculture, Agricultural Research
Service (USDA, ARS) (2013) USDA National
Nutrient Database for Standard Reference, Release 26.
Nutrient Data Laboratory Home Page, http://www.ars.
usda.gov/ba/bhnrc/ndl
Vardhini BV, Sujatha E, Rao SSM (2012) Influence of
brassinosteroids on metabolites of Raphanus sativus
L. J Phytol 4(2):45–47
Vargas R, Perez RM, Perez S, Zavala MA, Perez C (1999)
Antiurolithiatic activity of Raphanus sativus aqueous
extract on rats. J Ethnopharmacol 68(61–3):335–338
Visentin M, Tava A, Iori R, Palmieri S (1992) Isolation
and identification of trans-4-(methylthio)-3-butenyl
glucosinolate from radish roots (Raphanus sativus L.).
J Agric Food Chem 40(9):1687–1691
Wada K, Onda M, Matsubara H (1989) Amino acid
sequences of ferredoxin isoproteins from radish roots.
J Biochem 105(4):619–625
Wang LS, Sun XD, Cao Y, Wang L, Li FJ, Wang YF
(2010) Antioxidant and pro-oxidant properties of acyl-
ated pelargonidin derivatives extracted from red radish
(Raphanus sativus var. niger, Brassicaceae). Food
Chem Toxicol 48(10):2712–2718
Wu XL, Beecher GR, Holden JM, Haytowitz DB,
Gebhardt SE, Prior RL (2006) Concentrations of
anthocyanins in common foods in the United States
and estimation of normal consumption. J Agric Food
Chem 54(11):4069–4075
Wu SB, Wang F, Jin XB, Liu T, Wu LR, Wang W, Mei D,
Zheng Z, Zhu JY (2014) Two new natural products
from the roots of Raphanus sativus L. Adv Mat Res
884–885:548 (In Chinese)
Yamasaki M, Omi Y, Fujii N, Ozaki A, Nakama A,
Sakakibara Y, Suiko M, Nishiyama K (2009) Mustard
oil in “Shibori Daikon” a variety of Japanese radish,
selectively inhibits the proliferation of H-ras-
transformed 3Y1 cells. Biosci Biotechnol Biochem
73(10):2217–2221
Yang YW, Tai PY, Chen Y (2000) The origin of Raphanus
sativus based on the DNA sequences from different
organelles. HortScience 35(3):397
Raphanus raphanistrum subsp. sativus
Yang F, Lian GY, Yu B (2010) Synthesis of raphanuside,
an unusual oxathiane-fused thioglucoside isolated
from the seeds of Raphanus sativus L. Carbohydr Res
345(2):309–314
Yoshida Y, Takada N, Koda Y (2010) Isolation and identi-
fication of an anti-bolting compound, hexadecatrie-
noic acid monoglyceride, responsible for inhibition of
bolting and maintenance of the leaf rosette in radish
plants. Plant Cell Physiol 51(8):1341–1349
869
Zhang MF, Shen YQ (1996) Antidiarrheal and antiinflam-
matory effects of sinapine. Pharmcol Clin Chin
Materia Medica 12(1):29–31
Zhang YH, Yang ZH, Cao ZX (1983) Studies on stigma
pellicle glycoproteins of Raphanus sativus L. Acta Bot
Sinica 25(6):544–550
Zhang X, Liu HB, Jia JJ, Lv WH (2010) Two novel sulfur
compounds from the seeds of Raphanus sativus L. J
Asian Nat Prod Res 12(2):113–118