Content uploaded by Piero Picciarelli
Author content
All content in this area was uploaded by Piero Picciarelli on Dec 30, 2013
Content may be subject to copyright.
REVIEW
Globe artichoke as a functional food
Nello Ceccarelli •Maurizio Curadi •
Piero Picciarelli •Luca Martelloni •
Cristiana Sbrana •Manuela Giovannetti
Received: 18 June 2010 / Accepted: 29 June 2010 / Published online: 13 July 2010
ÓSpringer-Verlag 2010
Abstract Globe artichoke (Cynara cardunculus var.
scolymus L. Fiori) is a traditional component of the Med-
iterranean diet. Artichoke edible parts are one of the richest
dietary sources of polyphenols with high bioavailability,
and contain also high-quality inulin, fibres and minerals.
Moreover, pharmaceutical artichoke leaf extracts show
hypocholesterolemic and choleretic properties. Several
clinical studies have shown that the bioactive properties of
globe artichoke are due to the high content of polyphenolic
compounds in flower heads and leaves, in particular
hydroxycinnamates and flavonoids. The total antioxidant
capacity of artichoke flower heads is one of the highest
reported for vegetables, and is strictly related to their
polyphenolic content. Different agronomic variables, such
as mycorrhizal inoculation, may affect plant secondary
metabolism, and in particular the production of metabolites
with health-promoting activities. Our recent data showed
large increases of total phenolic content and antioxidant
activity in leaves and flower heads of mycorrhizal arti-
choke plants.
Keywords Artichoke Functional foods Polyphenols
Mycorrhizal symbiosis Antioxidant activity
Introduction
Globe artichoke (Cynara cardunculus var. scolymus
L. Fiori) is an ancient perennial plant species (Asteraceae)
native to the Mediterranean Basin and known since the first
century A.D. The Arabs played an important role in its
diffusion in the Southern Mediterranean area during the
Middle Ages [1]. Nowadays artichoke contributes signifi-
cantly to the Mediterranean agricultural economy, and Italy
is the main world producer. Globe artichoke is cultivated
for its large immature flower heads. The edible parts are the
tender inner bracts and the receptacle, commonly known as
‘‘heart’’ that constitute nearly 35–55% of the fresh weight
of the head.
In recent years, several epidemiological studies showed
that the consumption of fruits and vegetables, typical of the
Mediterranean diet, is associated with a decrease in the risk
of cancer and cardiovascular diseases [2]. Such health
benefits are mainly due to the high content of antioxidant
compounds (i.e. polyphenols, ascorbic acid, carotenoids) in
plant foods. Plant polyphenols are the most abundant
source of antioxidants in our diet [3]. Compared to other
vegetables, artichoke flower heads contain high levels of
total polyphenols [4] with hepatoprotective, hypocholes-
terolemic and antioxidant properties [5,6]. For these
characteristics, artichoke—a traditional component of the
Mediterranean diet—may be considered a functional food,
according to the definition of the European Commission on
Functional Food Science in Europe (FuFoSE) [7]. From a
nutritional point of view, artichoke is a plant food whose
natural components, beyond their basic nutritional value,
have positive effects on particular target functions, there-
fore reducing the risk of specific diseases. Artichoke has
also several non-food uses: the leaves, rich in polyphenols,
are employed by the pharmaceutical industry for the
N. Ceccarelli M. Curadi P. Picciarelli L. Martelloni
M. Giovannetti (&)
Department of Crop Plant Biology, University of Pisa,
Viale delle Piagge 23, 56124 Pisa, Italy
e-mail: mgiova@agr.unipi.it
C. Sbrana
Institute of Biology and Agrobiotechnology, CNR,
UOS Pisa, Via del Borghetto 80, 56124 Pisa, Italy
123
Mediterr J Nutr Metab (2010) 3:197–201
DOI 10.1007/s12349-010-0021-z
production of commercial extracts, while roots and flower
heads contain inulin, an important oligosaccharide used as
a prebiotic ingredient in functional foods [8]. Artichoke
industrial by-products (stems, outer bracts, leaves) repre-
sent about the 80% of the biomass and may be used as raw
material for the extraction of food additives and nutra-
ceuticals [6]. Moreover, given the large amount of ligno-
cellulosic biomass produced during its cultural cycle,
artichoke may be used as green forage for livestock and as
an energy crop [9].
Italian varieties and propagation
Both globe artichoke and cultivated cardoon (Cynara
cardunculus var. altilis DC) derive from the wild Cynara
cardunculus var. sylvestris Fiori. Globe artichoke is a
robust herbaceous perennial plant with vigorous growth,
medium tolerance to salinity and great adaptation to
Mediterranean climates. It has an autumn-spring growth,
from September (emergence) to July (seed maturity), and
efficiently uses rainfall as water supply. The commercial
production of artichoke is mainly based on the perennial
cultivation of vegetatively propagated clones. With this
objective in view, stems originating from buds on rhizomes
(known as ‘‘carducci’’) are traditionally used, although in
recent years both micropropagation and seed-planted
hybrids have been developed. Clonal propagation of arti-
choke has several advantages, in particular the mainte-
nance of uniformity and heterosis, but it poses problems for
the modern field rotation (an artichoke crop may last six or
more years, reaching the maximum productivity in the
third year), and for the build-up of pathogens and viruses.
Micropropagation techniques are used to obtain pathogen-
free plantlets of late varieties, also taking advantage of
hormonal treatments. Seed-propagated varieties have con-
verted this vegetable from a perennial into an annual crop,
more suitable for industrial agriculture, despite the loss of
precocity and heterosis. On the other hand, the traditional
diffusion in Italy of artichoke clonal propagation by means
of ‘‘carducci’’ has allowed the maintenance of a great
variety of local germplasm of high qualitative value, pre-
venting the loss of genetic diversity, and this is important
for a plant species which is not only a traditional crop, but
also a source of pharmaceutical compounds and a func-
tional food. From this point of view, the preservation of the
genetic diversity of artichoke germplasm as a natural
source of bioactive compounds is fundamental, both for
nutraceutic/pharmacologic applications, and for the selec-
tion of new artichoke varieties with enhanced polyphenolic
content.
Molecular studies showed that the varieties of globe
artichoke cultivated in Italy are highly heterogeneous and
this reflects their multiclonal composition, a direct conse-
quence of the limited selection criteria applied by farmers
[10], aimed at improving earliness and yield, based on
interclonal hybridization and intraclonal selection [11].
Commonly, four main artichoke varietal groups are dis-
tinguished in Italy (‘‘Spinosi’’, ‘‘Violetti’’, ‘‘Catanesi’’ and
‘‘Romaneschi’’), but most local germplasm is not fully
included in these categories, showing a great range of
variation both for productive characters, in particular
flower head parameters (dimension, shape, weight, n°per
plant, earliness, etc.), and for qualitative-quantitative
phenolic profile [12]. On the basis of the phenological
cycle, reflowering (=early) varieties (i.e. ‘‘Spinoso Sardo’’,
‘‘Catanese’’, ‘‘Violetto di Sicilia’’) and late varieties
(i.e. ‘‘Romanesco’’, ‘‘Violetto di Toscana’’, ‘‘Terom’’) are
classified.
Artichoke production in Italy
Artichoke iswidely distributed all over the world (126,429 ha)
with a production of 1,386,848 t and average yields of 109
q/ha, although it is mainly concentrated in the Mediterra-
nean regions [13]. Italy is the leading producer (40% of the
global production) with 50,699 ha and 510,141 t per year,
followed by Spain (16,800 ha), France (9,475 ha) and
Egypt (3,800 ha) [14]. In Southern Italy, artichoke
is mainly cultivated in Puglia (16,930 ha), Sicilia
(14,800 ha) and Sardegna (13,630 ha) that produce early
varieties, while in Central Italy late varieties are cultivated
in Campania (2,012 ha), Lazio (1,058 ha) and Toscana
(548 ha) [14]. Fresh flower heads are the main prod-
uct, but consistent amounts are also used for industrial
transformation.
Due to the high production costs and the low market
prices caused by the competition of other Mediterranean
countries (mainly Spain and Egypt), Italian artichoke needs
to be promoted through marketing strategies aimed at
stressing qualitative features that may positively influence
consumers’ choice. Even though traditional qualitative
parameters (organoleptic, commercial) are still used as
reference for fresh vegetables, in recent years consumers
have shown great interest in other qualitative characteris-
tics, such as safety, nutritional value and nutraceutical
content.
Quality of artichoke: safety, nutritional value
and nutraceutical content
Artichoke is a genetically robust plant with a marked tol-
erance to pathogens and aphids: this is probably due to the
high content of sesquiterpenes with antifeedant activity
198 Mediterr J Nutr Metab (2010) 3:197–201
123
(such as cynaropicrin, which is also responsible for the
typical bitter taste of artichoke). In Italy, local artichoke
ecotypes are highly adapted to the Mediterranean climate:
they show a great tolerance to biotic and abiotic stresses,
allowing the cultivation without the aid of chemical
treatments.
Artichoke flower heads contain about 15–20% of dry
matter, with a remarkable nutritional value (6.8% carbo-
hydrates, 2.9% nitrogen substances). Vitamin C content is
high: 10 mg/100 g f.w. [15]. Compared to other vegeta-
bles, artichoke flower heads are particulary rich in inulin
(19–36% d.w.), with the highest degree of polymerisation
known in plants [6]. Artichoke flower heads represent also
a rich source of minerals, showing in particular K and Ca
contents of 360 and 50 mg/100 g f.w., respectively [16].
Artichoke flower heads, together with blueberries and
soybeans, are among the richest sources of dietary phenolic
antioxidants, showing a total antioxidant capacity (TAC) of
more than 9,000 lmol of Trolox equivalents/100 g f.w.
[17].
From a nutritional point of view, the high levels of
bioactive polyphenols (caffeoylquinic acids and flavo-
noids) in the inner bracts and receptacles represent an
added value for artichoke flower heads. Polyphenolic
content in artichoke edible parts is variable: values ranging
from 4.8 mg/g f.w. in var. ‘‘Spinoso Sardo’’ [18]to
29.8 mg/g f.w. in var. ‘‘Violetto di Toscana’’ [16] are
reported in the literature. In a recent study, carried out at
the Department of Crop Plant Biology of University of
Pisa, the total phenolic content (TPC, expressed as lgof
chlorogenic acid equivalents/mg f.w.) in the edible parts
of early and late artichoke varieties at the harvest stage
ranged from 10.4 to 18.6% d.w. (corresponding to 7.3 and
13 mg/g f.w., respectively) [19].
The qualitative and quantitative variability of the phe-
nolic complement in artichoke flower heads of different
varieties depends on their genetic diversity, as well as on
their physiological stage of development (harvest time) and
on climatic conditions during plant growth [12,20]. In a
recent molecular study, Italian artichoke genotypes have
shown highly different genetic backgrounds and their
phenolic content appeared strongly influenced by genetic
factors [10]. Other authors have reported, for American
varieties (‘‘Green Globe’’), lower TPC values compared to
European ones [21]: such data may lead to hypothesise that
Mediterranean varieties display a higher polyphenolic
content due to positive interactions between genetic and
environmental factors.
In recent years, evidence of the healthy functions of
polyphenols assumed with the diet have led to investigate
their biosynthesis in plants, and several key enzymes of the
phenylpropanoid pathway, as well as their encoding genes,
have been studied in different plant species. Overexpression
of phenylalanine ammonia lyase (PAL) has been shown
to increase polyphenol accumulation in tobacco [22]. In globe
artichoke, two acyltransferases, HCT (hydroxycinnamoyl-
CoA: shikimate/quinate hydroxycinnamoyltransferase) and
HQT (hydroxycinnamoyl-CoA quinate: hydroxycinnamoyl-
transferase), involved in the synthesis of caffeoylquinic
acids, have been isolated and characterised [23,24].
Enhancement of phenolic complement is presently pursued
in various crops by genetic engineering [25].
Biological activities of artichoke polyphenols
The bioactivity displayed by artichoke flower heads and
leaves is strictly related to their polyphenolic content.
Artichoke leaves contain a very high level of total pheno-
lics, and this justifies their wide use in phytopharmaceutical
applications [12]. Artichoke leaf extracts are currently
commercialised as drugs mainly for the treatment of liver
diseases. In various pharmacological studies, artichoke leaf
extracts have shown a wide range of effects, including
choleretic, hypocholesterolemic and antioxidant activities
[6]. These bioactivities cannot be ascribed to a single
molecule, but to several compounds, as a result of additive
or synergistic interactions. The diuretic and hepatostimu-
lant effects of artichoke leaves have been known since
ancient times, but only in the last decades the bioactive
constituents have been identified.
Chlorogenic acid (5-O-caffeoylquinic acid), 1,5- and
3,4-di-O-caffeoylquinic acids and cynarine are the pre-
dominant compounds among hydroxycinnamates, while
the main flavonoids are apigenin and luteolin, and their
glycosides [12,26]. These phenolics display a marked
scavenging activity against reactive oxygen species (ROS)
and free radicals, acting as a protective pool against oxi-
dative damage to biological molecules, such as proteins,
lipids and DNA [27,28].
The pharmacologic properties of artichoke leaf extracts
are well documented in several in vitro and in vivo studies.
The choleretic activity (increased biliary flux with elimi-
nation of cholesterol) of both chlorogenic acid and cynar-
ine has been demonstrated in several clinical trials [29,30],
and artichoke leaf extracts are believed to be effective in
the therapeutic treatment of dyspeptic syndromes and
gastric diseases. Artichoke leaf extracts also inhibit the
hepatic biosynthesis of cholesterol. This effect is due to
luteolin, which modulates the HMG-CoA reductase activ-
ity (the key enzyme in the cholesterol biosynthesis path-
way) by inhibition mechanisms [31,32]. Moreover,
chlorogenic acid and luteolin may prevent atherosclerosis
inhibiting low-density lipoproteins (LDL) oxidation [33].
Therefore, artichoke leaf extracts show hypocholestero-
lemic activity, due to two parallel mechanisms: reduction
Mediterr J Nutr Metab (2010) 3:197–201 199
123
of cholesterol biosynthesis and inhibition of LDL oxidation
[34,35]. Artichoke extracts are well tolerated, and may be
useful for the preventive treatments of mild hypercholes-
terolemia.
In recent years, the increasing demand for functional
foods led to a great interest in natural compounds with
antioxidant properties. Since the antioxidant content is
considered an important quality parameter for vegetables,
and a key factor for market development, the acquisition
and diffusion of scientific information on the nutritional
value and health effects of artichoke may influence the
consumers’ food choices.
Whereas past researches concentrated on the phenolic
content of artichoke leaves, used by the pharmaceutical
industry, more recently there is a growing interest in the
phenolic complement of fresh edible parts, also in relation
to different genotypes. Chlorogenic acid is by far the
quantitatively predominant polyphenol in artichoke flower
heads, ranging from 1.3 to 2.4 mg/g f.w. [19]; among
flavonoids, apigenin-7-O-glucuronide is the most repre-
sented [12]. Dietary polyphenols may exert their biological
activity only when they are present in the cells of target
tissues at the right concentration. Therefore, their effective
bioactivity depends on the amount assumed with the diet as
well as on the rate of absorption in the gut and on
metabolism. Studies on the absorption and metabolism of
plant bioactive compounds in humans, often overlooked in
the past, are now considered fundamental for the inter-
pretation of in vivo effects. In recent clinic trials, high
bioavailability of chlorogenic acid after oral administration
of both artichoke leaf extracts [36] and cooked edible parts
[37] was shown. Chlorogenic acid is rapidly adsorbed in
the human gastrointestinal tract or hydrolysed to caffeic
acid, which also exerts antioxidant activity.
Nutraceutical content of mycorrhizal artichoke
Arbuscular mycorrhizas are one of the most widespread
mutualistic symbioses, established between the roots of
most land plants and fungi of the phylum Glomeromycota
(arbuscular mycorrhizal fungi, AMF). AMF increase plant
nutrition and water uptake by means of an extensive
mycelial network spreading from colonised roots into the
soil [38,39]. AMF contribute to the maintenance of soil
fertility and may be considered ‘‘natural biofertilizers’’,
whose importance is fundamental in sustainable agroeco-
systems. In the host plant this association determines
higher growth rates, increased resistance to root pathogens
and abiotic stresses (water deficit and salinity), and
increased biosynthesis of antioxidant compounds [40].
Arbuscular mycorrhizas are found in about 80% of plant
species, including cereals, legumes, fruit plants and
vegetables. Present day trend towards a more sustainable
agriculture, with low chemical inputs, leads to consider
mycorrhizal inoculation with suitable species of AMF a
potential biofertilization strategy, in order to enhance plant
growth, yield and quality.
A recent work investigated the effects of mycorrhizal
inoculation with the AMF species Glomus mosseae and
Glomus intraradices (alone or in mixture) on the poly-
phenolic content and the antioxidant properties of arti-
choke. Mycorrhizal symbiosis enhanced phenolic contents
in both leaves and flower heads and the most significant
effect was observed in plants inoculated with the mixture
of both fungal species. Interestingly, mycorrhizal inocula-
tion responses persisted for 2 years after field transplant.
Flower heads of such plants showed large TPC increases
(47 and 55% in the first and second year in the field,
respectively) and ARP enhancements (52 and 30% in the
first and second year in the field, respectively) in compar-
ison with control plants. Moreover, the mean weight of main
flower heads was about 92 and 70% higher (first and second
year in the field, respectively) compared to control plants [41].
Our data allowed us to demonstrate that beneficial symbionts,
such as AMF, represent an environmentally friendly and
efficient strategy to enhance plant biosynthesis of secondary
metabolites with health-promoting activities.
Conflict of interest None.
References
1. Sonnante G, Pignone D, Hammer K (2007) The domestication of
artichoke and cardoon: from Roman times to the genomic age.
Ann Bot 100:1095–1100
2. Williamson G, Manach C (2005) Bioavailability and bioefficacy
of polyphenols in humans. II. Review of 93 intervention studies.
Am J Clin Nutr 81:243S–255S
3. Manach C, Scalbert A, Morand C, Remesy C, Jimenez L (2004)
Polyphenols: food sources and bioavailability. Am J Clin Nutr
79:724–747
4. Brat P, George
`S, Bellamy A, Du Chaffaut L, Scalbert A, Mennen
L, Arnault N, Amiot MJ (2006) Daily polyphenol intake in
France from fruit and vegetables. J Nutr 136:2368–2373
5. Shutz K, Kammerer D, Carle R, Schieber A (2004) Identification
and quantification of caffeoylquinic acids and flavonoids from
artichoke (Cynara scolymus L.) heads, juice and pomace by
HPLC-DAD-ESI/MS
n
. J Agric Food Chem 52:4090–4096
6. Lattanzio V, Kroon PA, Linsalata V, Cardinali A (2009) Globe
artichoke: a functional food and source of nutraceutical ingredi-
ents. J Funct Foods 1:131–144
7. Roberfroid MB (2000) A European consensus of scientific con-
cepts of functional foods. Nutrition 16:689–691
8. Raccuia SA, Melilli MG (2004) Cynara cardunculus L., a
potential source of inulin in the Mediterranean environment:
screening of genetic variability. Aust J Agric Res 55:693–698
9. Megı
`as MD, Hernandez F, Madrid J, Martinez-Teruel A (2002)
Feeding value, in vitro digestibility and in vivo gas production of
different by-products for ruminant nutrition. J Sci Food Agric
82:567–572
200 Mediterr J Nutr Metab (2010) 3:197–201
123
10. Moglia A, Lanteri S, Comino C, Acquadro A, Devos R, Beek-
wilder J (2008) Stress-induced biosynthesis of dicaffeoylquinic
acids in globe artichoke. J Sci Food Agric 56:864
11. Lanteri S, Di Leo I, Ledda L, Mameli MG, Portis E (2001) RAPD
variation within and among populations of globe artichoke cul-
tivar ‘‘Spinoso sardo’’. Plant Breed 120:243–246
12. Lombardo S, Pandino G, Mauromicale G, Knodler M, Carle R,
Schieber A (2010) Influence of genotype, harvest time and
plant part on polyphenolic composition of globe artichoke
[Cynara cardunculus L. var. scolymus (L.) Fiori]. Food Chem
119:1175–1181
13. FAO Statistical Database (2008) http://faostat.fao.org/
14. ISTAT (2009) http://www.istat.it/
15. Gil-Izquierdo A, Gil MI, Conesa MA, Ferreres F (2001) The
effects of storage temperatures on vitamin C and phenolic content
of artichoke (Cynara scolymus L.) heads. Innov Food Sci Emerg
Technol 2:199–202
16. Romani A, Pinelli P, Cantini C, Cimato A, Heimler D (2006)
Characterization of Violetto di Toscana a typical Italian variety of
artichoke (Cynara scolymus L.). Food Chem 95:221–225
17. Pennington JAT, Fisher RA (2009) Classification of fruits and
vegetables. J Food Compost Anal 22S:S23–S31
18. Alamanni MC, Cossu M (2003) Antioxidant activity of the
extracts of the edible part of artichoke (Cynara scolymus L.) var.
spinoso sardo. Ital J Food Sci 15:187–195
19. Curadi M, Picciarelli P, Lorenzi R, Graifenberg A, Ceccarelli N
(2005) Antioxidant activity and phenolic compounds in the edible
parts of early and late Italian artichoke (Cynara scolymus) vari-
eties. Ital J Food Sci 17:33–43
20. Lattanzio V, Cicco N, Linsalata V (2005) Antioxidant activities
of artichoke phenolics. Acta Hortic 681:421–428
21. Wang M, Simon JE, Aviles IF, He K, Zheng Q, Tadmor Y (2003)
Analysis of antioxidative phenolic compounds in artichoke
(Cynara scolymus L.). J Agric Food Chem 51:601–608
22. Chang J, Luo J, He G (2009) Regulation of polyphenols accu-
mulation by combined overexpression/silencing key enzymes
of phenylpropanoid pathway. Acta Biochim Biophys Sin 41:
123–130
23. Comino C, Lanteri S, Portis E, Acquadro A, Romani A, Hehn A,
Larbat R, Bourgaud F (2007) Isolation and functional charac-
terization of a cDNA coding a hydroxycinnamoyltransferase
involved in phenylpropanoid biosynthesis in Cynara cardunculus
L. BMC Plant Biol 7:14
24. Comino C, Hehn A, Moglia A, Menin B, Bourgaud F, Lanteri S,
Portis E (2009) The isolation and mapping of a novel hydro-
xycinnamoyltransferase in the globe artichoke chlorogenic acid
pathway. BMC Plant Biol 9:30
25. Niggeweg R, Michael A, Martin C (2004) Engineering plants
with increased levels of the antioxidant chlorogenic acid. Nat
Biotechnol 22:746–754
26. Pandino G, Courts FL, Lombardo S, Mauromicale G, Williamson
G (2010) Caffeoylquinic acids and flavonoids in the immature
inflorescence of globe artichoke, wild cardoon and cultivated
cardoon. J Agric Food Chem 58:1026–1031
27. Kono Y, Kobayashi K, Tagawa S, Adachi K, Ueda A, Sawa Y,
Shibata H (1997) Antioxidant activity of polyphenolic in diets:
rate constant of reactions of chlorogenic acid and caffeic acid
with reactive species of oxygen and nitrogen. Biochem Biophys
Acta Gen Sub 1335:335
28. Pavlica S, Gebhardt R (2005) Protective effects of ellagic and
chlorogenic acids against oxidative stress in PC12 cells. Free
Radic Res 39:1377–1390
29. Kirchhoff R, Beckers C, Kirchhoff GM, Trinczek-Gartner H,
Petrowicz O, Reimann HJ (1994) Increase in choleresis by means
of artichoke extract. Phytomedicine 1:107–115
30. Fintelmann V (1996) Therapeutic profile and mechanisms of action
of artichoke leaf extracts: hypolipemic, antioxidant, hepatoprotec-
tive and choleretic properties. Phytomedicine Suppl 1:50
31. Kraft K (1997) Artichoke leaf extracts: recent findings reflecting
effects on lipid metabolism, liver and gastrointestinal tracts.
Phytomedicine 4:369
32. Gebhardt R (2002) Inhibition of cholesterol biosynthesis in
HepG2 cells by artichoke extracts is reinforced by glucosidase
pretreatment. Phytother Res 16:368–372
33. Brown JE, Rice-Evans CA (1998) Luteolin-rich artichoke
extracts protect low density lipoprotein from oxidation in vitro.
Free Radic Res 29:247–255
34. Gebhardt R (1997) Antioxidative and protective properties of
extracts from leaves of the artichoke (Cynara scolymus L.)
against hydroperoxide-induced oxidative stress in cultured rat
hepatocytes. Toxicol Appl Pharmacol 144:279–286
35. Bundy R, Walker AF, Middleton RW, Wallis C, Simpson HCR
(2008) Artichoke leaf extract (Cynara scolymus L.) reduces
plasma cholesterol in otherwise healthy hypercholesterolemic
adults: a randomized, double blind placebo controlled trial.
Phytomedicine 15:668–675
36. Wittemer SM, Ploch M, Windeck T, Muller SC, Drewelow B,
Derendorf H, Veit M (2005) Bioavailability and pharmacoki-
netics of caffeoylquinic acids and flavonoids after oral adminis-
tration of artichoke leaf extracts in humans. Phytomedicine
12:28–38
37. Azzini E, Bugianesi R, Romano F, Di Venere D, Miccadei S,
Durazzo AM, Foddai MS, Catasta G, Linsalata V, Maiani G
(2007) Absorption and metabolism of bioactive molecules after
oral consumption of cooked edible heads of Cynara scolymusL
(cv. Violetto di Provenza) in human subjects: a pilot study. Br J
Nutr 97:963–969
38. Giovannetti M, Avio L, Fortuna P, Pellegrino E, Sbrana C, Strani
P (2006) At the root of the wood wide web: self recognition and
nonself incompatibility in mycorrhizal networks. Plant Signal
Behav 1:1–5
39. Giovannetti M, Avio L (2002) Biotechnology of arbuscular
mycorrhizas. In: Khachatourians GG, Arora Dilip K (eds)
Applied mycology and biotechnology, vol 2. Agriculture and
Food Production. Elsevier Science BV, Amsterdam, NL
40. Fester T, House B, Schmidt D, Hafmann K, Schmidt J, Wray V,
Hause G, Strack D (2002) Occurence and localization of apoca-
rotenoids in arbuscular mycorrhizal plant roots. Plant Cell
Physiol 43:256–265
41. Ceccarelli N, Curadi M, Picciarelli P, Martelloni L, Sbrana C,
Giovannetti M (2010) Mycorrhizal inoculation responses persist
two years after transplant and increase phenolic content and
antioxidant properties of artichoke leaves and flower heads. Plant
Soil. doi:10.1007/s11104-010-0417-z
Mediterr J Nutr Metab (2010) 3:197–201 201
123