Environmental eco-physiology and economical potential of the halophyte Crithmum maritimum L. (Apiaceae)

Abstract

The present contribution reviews information on Crithmum maritimum L., a facultative halophyte belonging to the Apiaceae family and typical of coastal ecosystems. It grows wild on maritime rocks, piers, breakwaters and sandy beaches along the Mediterranean, Pacific and Atlantic coasts. Its propagation by germination, vegetative multiplication and the in vitro culture techniques is quite easy. Salinities exceeding 50 mM NaCl inhibit seed germination but seem without impact on seed viability. At the vegetative stage, growth was stimulated by low salinity but was markedly reduced at high levels of salt without any symptoms of toxicity. C. maritimum L. has been largely used for nutritional and medicinal purposes. The plant is also edible and it is consumed in the traditional diet of the first European farmers, as a potent source of minerals, vitamin C, essential oils and other biomolecules. The fruit of C. maritimum L. is rich in lipids (about 44% on dry weight basis) with oleic acid as major component (78.6% of the total fatty acids). All these features make this species one of the most promising halophytes in the context of biosaline agriculture.

Full text

Journal of Medicinal Plants Research Vol. 5(16), pp. 3564-3571, 18 August, 2011
Available online at http://www.academicjournals.org/JMPR
ISSN 1996-0875 ©2011 Academic Journals
Review
Environmental eco-physiology and economical
potential of the halophyte Crithmum maritimum L.
(Apiaceae)
Abdallah Atia*, Zouhaier Barhoumi, Rabhi Mokded, Chedly Abdelly and Abderrazak Smaoui
Laboratoire des Plantes Extrêmophiles, Centre de Biotechnologies, Technopole de Borj Cedria, BP 901, Hammam-Lif,
2050, Tunisia.
Accepted 6 July, 2011
The present contribution reviews information on Crithmum maritimum L., a facultative halophyte
belonging to the Apiaceae family and typical of coastal ecosystems. It grows wild on maritime rocks,
piers, breakwaters and sandy beaches along the Mediterranean, Pacific and Atlantic coasts. Its
propagation by germination, vegetative multiplication and the in vitro culture techniques is quite easy.
Salinities exceeding 50 mM NaCl inhibit seed germination but seem without impact on seed viability. At
the vegetative stage, growth was stimulated by low salinity but was markedly reduced at high levels of
salt without any symptoms of toxicity. C. maritimum L. has been largely used for nutritional and
medicinal purposes. The plant is also edible and it is consumed in the traditional diet of the first
European farmers, as a potent source of minerals, vitamin C, essential oils and other biomolecules. The
fruit of C. maritimum L. is rich in lipids (about 44% on dry weight basis) with oleic acid as major
component (78.6% of the total fatty acids). All these features make this species one of the most
promising halophytes in the context of biosaline agriculture.
Key words: Crithmum maritimum, salinity, nutritional and medicinal uses, biosaline agriculture.
INTRODUCTION
Crithmum maritimum L. (Apiaceae) is a halophyte also
known as crest marine, marine fennel, sea fennel,
sampier and rock samphire (Atia et al., 2009a). It is
typical of rocky coastal ecosystems, since it grows wild
on maritime rocks, piers and breakwaters and sandy
beaches. This aromatic plant grows wild in rock crevices,
rocky shores and shingle beaches along the
Mediterranean and Black sea coasts, as well as along the
Atlantic coast of Portugal and of south and south-west
England, Wales and Southern Ireland. This plant also
occurs along the coasts of other countries (e.g. Canada)
as a naturalized species (Ozcan, 2000; Ben Amor et al.,
*Corresponding author. E-mail: atbdllh@yahoo.fr. Tel: (+216) 79
412 848. Fax: (+216) 79 412 638.
ABBREVIATIONS: ABA, abscisic acid; ABTS, 2,2'-azino-bis(3-
ethylbenzthiazoline-6-sulphonic acid); BA, 6-Benzyladenine;
B5, gamborg medium; GA
3,
gibberellic acid 3; IBA, indole-3-
butyric acid; MS, murashige and skoog medium; MDA,
malondialdehyde.
2005; Cornara et al., 2009). C. maritimum L. shows
substantial economical and medicinal potentials: it is
edible aromatic and has a powerful scent. Its organs
(roots, leaves and fruits) are rich in several bioactive
substances that could be used as aromatic, medicinal,
antimicrobial and insecticide (Atia et al., 2009a; Meot-
Duros and Magné 2009; Meot-Duros et al., 2010). In
temperate climates, the plant is used for ornamental
decoration in rock gardens along the sea (Franke, 1981).
It is also cultivated in many areas across Europe for
several economic and industrial purposes (Franke, 1981;
Atia et al., 2010a; Meot-Duros et al., 2008).
In the recent years, products of halophytes are more
and more produced and sold in the markets through the
world (Geissler et al., 2009; Koyro and Lieth, 2008; Koyro
et al., 2008). The benefit of herbal drugs has been
considered since ancient times (Grabley and Thiericke,
1999; Cornara et al., 2009), and even today many
halophytes are investigated for the development of new
phytocompounds. The demand from the medicinal and
cosmetic industry for essential oils or other products of C.
maritimum is rapidly increasing (Grigoriadou and

Maloupa, 2008). The objective of this review is to go over
and to update our knowledge about the medicinal
halophyte C. maritimum with a particular emphasis on its
eco-physiological responses to saline environment, its
economic importance and its potential as a promising
candidate for bio-saline agriculture.
CLASSIFICATION
Taxonomy
Kingdom: Plantae
Subkingdom: Tracheobionta
Superdivision: Spermatophyta
Division: Magnoliophyta
Class Magnoliopsida
Subclass: Rosidae
Order: Apiales
Family: Apiaceae
Genus: Crithmum
Species: Crithmum maritimum L.
Crithmum: from Greek krithe: barley, from resemblance
of fruit to barleycorn.
Maritimum: of the sea.
MORPHOLOGICAL DESCRIPTION
C. maritimum L. (Apiaceae) is a highly branched
perennial herb of up to 30 to 60 cm in height (Cornara et
al., 2009). The root is a strong, thick and gnarled (Franke,
1981). The leaves are fleshy and succulent Figure 1.
They extend radially forming a rosette. They have a
sheath-like base with the short petiole ending in a pinnate
compound blade, which is usually divided into 3 leaflets,
each again pinnate. The leaflets are 2 to 5 cm long, 0.6
cm wide and linear to lanceolate with a conical, some-
times spiny tip. From the end of July to mid-August, a
stalk up to 30 cm high with 2 or 3 smaller leaves
develops from the terminal bud and ends in a compound
umbel with 10 to 20 rays, bearing an involucre and
involucels, each one consisting of several leaflets.
The species flowers between June and September and
the fruit begins to mature in November-December. The
flowers are of yellowish or greenish-white colour and
produce ovoid fruits which consist of two mericraps each
of them subdivided in 5 ribs. The Fruit are 5 to 6 mm
long, 1.5 to 2.5 mm large, ovoid-oblong, not compressed,
corky and olive-green to purple (Figure 1). At maturation
stage, in each groove of the spongy mesocarp, several
brown lines are visible. They represent the vittae. The
carpophore is also present (Franke, 1981; Atia et al.,
2010b).
Cultivation and multiplication aptitude
This species produces high number of viable fruits that
Atia et al. 3565
permit their multiplication by germination without any
inconvenience. The germination of C. maritimum fruit has
been reported to be maximal in distilled water (Atia et al.,
2009a). The reed light, the nitrate and other nitrogen
compounds significantly promote and accelerate
germination (Atia et al., 2009b).
Although C. maritimum provide a limited number of
cuttings, the propagation by softwood cuttings is also
possible without any inconvenient. When the cuttings
produce sufficient roots, they can be easily transplanted
to the main field.
In vitro propagation is a useful technique for mass
multiplication and germplasm conservation of any plant
species (Kavitha et al., 2010). The multiplication of the C.
maritimum L. via in vitro culture technique has been
recently reported (Grigoriadou and Maloupa, 2008).
Different culture media have been used for in vitro culture
of other species of the family of Apiaceae (Hirai et al.,
1997). Shoot production of C. maritimum was significantly
stimulated when shoot tip explants were cultured in MS
medium. MS seems to be the most effective of the basal
media tested for in vitro cultivation of C. maritimum as it
leads to a significantly increase of number of new
microshoots produced / explant (3.4) and enhances shoot
height (3.1 cm).
According to Grigoriadou and Maloupa (2008), the B5
medium favours rooting (92.5% of the microshoots
develop roots) with and an average of 5.8 roots/explant
and 0.6 cm length. When added at 2.5 to 10 µM, BA
significantly increased shoot proliferation. Further, the
combination of different IBA concentrations with 2.5 µM
BA significantly improved the rooted microshoots, the
number of roots and root length.
ECO-PHYSIOLOGICAL RESPONSES TO SALT
STRESS
Germination stage
Despite C. maritimum L. usually grows in the vicinity of
seawater, salinities exceeding 50 mM NaCl were found to
inhibit its germination (Atia et al., 2006; Atia et al.,
2009a). In the natural conditions of C. maritimum L., fruits
are continuously exposed to various ions including; Na
+
,
Mg
2+
, Ca
2+
, Cl
-
, and SO
4
2-
. In a recent report, we show
that the salt-induced inhibition of seed germination was
salt-specific and could be classified in the following
decreasing order: MgCl
2
, MgSO
4
, Na
2
SO
4
, NaCl.
Magnesium salts, that is, MgCl
2
and MgSO
4
, restrict
germination via their osmotic and ionic effects. At verylow
osmotic potentials, Mg
2+
exerted a strong toxic effect that
may be explained by the high loss of nutrients from
seeds, especially phosphorus, nitrate, sulphate and
calcium.
Sodium salts, that is, Na
2
SO
4
and NaCl, adversely
affected germination mainly via an osmotic effect, since
high germination recovery could be observed after seed

3566 J. Med. Plant. Res.
C
D
E
F
G
F
A
B
H
Figure 1. General aspect of C. maritimum: (A) Aspect of cultivated plant, (B) detailed view of leaves. Note the
succulence aspect, (C) inflorescence aspect at the flowering stage, (D) inflorescence view after fruit formation,
(E) inflorescence aspect after fruit maturation, (F) and (G) detailed view of the fruit during prematuration stage,
(H) details view of fruit at maturation.
transfer in distilled water. This suggests that in natural
conditions, the plant produces seed banks consisting in
seeds that remain viable and germinate after the winter
rains, so that the plant can successfully establish (Atia et
al., 2011).
A useful approach to overcome the salt-induced seed
dormancy observed in halophytes consists in the exo-
genous application of germination-promoting substances.
In this way, nitrate, ammonium, and GA
3
proved to
significantly enhance seed germination of C. maritimum
L. under salinities (Atia et al., 2009a). Interestingly, red
light application was also efficient for seed germination
induction under salinity (Atia et al., 2009c). It seems
that salt inhibits germination partly by increasing ABA
content in seeds. This was confirmed by the fact that the
germination was inhibited by exogenous ABA addition in
the imbibition medium. This inhibition was alleviated by
nitrate (Figure 2). Fluridone, an inhibitor of ABA
synthesis, alleviated the salt-induced restriction of
germination too (Atia et al., 2009b).
In C. maritimum L. fruit, NaCl is mainly accumulated in
the external envelopes, that is, the spongy coat, the
secretory envelope and the endocarp layer (Atia et al.,
2010b). This phenomenon is of vital eco-physiological
significance for this halophyte, since it preserves the
embryo viability even if the salinity increases in the

Atia et al. 3567
MgCl
2
> MgSO
4
>
Seeds
Spongy coat
Reserve mobilisation
and embryo growth
Germination
Rainy water
ABA
Na
2
SO
4
>NaCl
Loss of viability
Lessivage of salt
Dispersion and
Protection
Chemicals: Flu,
nitrate, a
mmonium
and GAs
Salinity
Figure 2. A working model explaining the eco-physiological responses of C. maritimum to salinity during
germination stage.
medium (Figure 3). Upon salt leaching by winter rains,
water imbibition starts and seed germination can take
place allowing C. maritimum L. to establish in the saline
biotopes.
Vegetative stage
Several authors showed that C. maritimum L. is a
facultative halophyte (Ben Hamed et al., 2004; Ben Amor
et al., 2005). According to Ben Hamed et al. (2007), leaf
growth was stimulated by 50 mM NaCl, unaffected at 100
mM NaCl and was significantly decreased at 300 mM
NaCl but without any toxicity symptoms. Root growth was
significantly reduced at 100 and 300 mM NaCl (Ben
Hamed et al., 2007) Table 1.
In C. maritimum L., accumulating high amounts of Na
+
and Cl
-
in leaves had no severe impact on their water
status, which is indicative of includer behaviour (Ben
Hamed et al., 2004; Ben Amor et al., 2005, 2006). By
opposition to excluders halophyte which secrete the salt
by specific leaf structure like glands or trichomes,
includer halophytes sequester these toxic ions in their
vacuoles. Ben Amor et al. (2005) showed that salt
treatment reaching 200 mM NaCl did not induce
membrane lipid peroxidation since MDA values in both
roots and shoots remained close to those control. This
was concomitant with the stimulation of activities of the
protective antioxidant enzymes, namely superoxide
dismutase (SOD), catalase and peroxidase. Thus, the
antioxidative system was efficient to protect the plant
tissues against the toxic ions like Na+ and Cl- (Ben
Hamed et al., 2007) Table 1.
Salt tolerance aptitude of C. maritimum L. was also
attributed to its ability to maintain potassium supply, a
convenient water supply and/or highly capacity to
conserve tissue hydratation and to exhibit an efficient
antioxidant system (Ben hamed et al., 2004). This
performance is also related to a set of morphological
adaptations including leaf succulence, abundance of
palisade parenchyma, and aquifer parenchyma, a thick
cuticle layer and a low number of stomata avoiding water
loss.
ETHNOBOTANICAL AND MEDICINAL USES
C. maritimum L. has been largely used for nutritional and
medicinal purposes. The plant is edible; it was consumed
in the traditional diet of the first European farmers, as it is
a significant source of minerals. It was cultivated in
gardens and was sold on London streets as 'Crest
Marine' (Guil-Guerrero et al., 1998). The use of Samphire
as a condiment and pickle or as an ingredient in a salad

3568 J. Med. Plant. Res.
Quinic acid
Diosmin
Hesperidin Chlorogenic acid
Falcarindiol p-Toluic acid
Figure 3. Chemical structure of some biological active compounds found in C. maritimum L.
is well known (Atia et al., 2006; Meot-Duros et al., 2009).
For instance, in many European countries, the leaves are
washed, cut into small pieces and prepared for salads by
mixed juice and olive oil. In British Isles, the leaves were
formerly pickled and kept like capers in vinegar. Rock
Samphire Hash is a traditional British recipe. This was
prepared by mixing stems and leaves of C. maritimum L.
with a pickled cucumber and caper which cooked in
stock. Then this was bound with an egg yolk
(http://www.celtnet.org.uk/recipes/ancient/wild-food
entry.phpterm). In a Greek legend, it is even mentioned
as a vegetable served to Theseus by Hekate.
Cornara et al. (2009) reported that C. maritimum L. is
used in folk medicine as appetizer, tonic, carminative,
diuretic and vermifuge. Sailors used to consume food
preparations based on C. maritimum L. or eat leaves
asprotection against scurvy (Cunsolo et al., 1993). When
going on fishing trips, sailors took fresh leaves with them.
But on longer voyages the leaves were apparently kept
pickled in vinegar for better preservation. In Italy, the
decoction of shoots harvested before fructification were
used against inflammations of the urinary tract and
prostate and colics. It has tonic and purgative action
while the infusion of leaves has been largely used for the
digestive diseases and for renal therapy (Franke, 1981:
Cunsolo et al., 1993; Guil-Guerrero and Rodriguez-

Atia et al. 3569
Table 1. Summary of the eco-physiological salt-responses during the vegetative stage of C. maritimum L.
Physiological parameter Low salinity Moderate salinity High salinity References
Growth Stimulated Not affected Reduced Ben Amor et al. (2005)
Tissue hydratation Not affected Not affected reduced Grigoriadou and Maloupa (2008)
Toxic ion accumulation Accumulated Accumulated Highly accumulated Ben Hamed et al. (2004)
Potassium uptake Maintained Maintained Maintained Ben Amor et al. (2005)
Photosynthetic activity Maintained Maintained Maintained Ben Hamed et al. (2004)
Antioxidant systems Stimulated Stimulated Reduced
Ben Amor et al. (2005)
Ben Hamed et al. (2007)
Toxicity or tolerance Tolerance Tolerance Tolerance Ben Amor et al. (2006)
Table 2. Geographical variability in the major volatile compounds from C. maritimum L.
(adapted from Pateira et al. 1999).
Compounds
Portugal France Italy
Sabinene 24.4 - 0.7
α-Pinen 0.2 - 0.1
Cis-ß-Ocimen 3.9 - -
y-Terpinene 35.0 1% 22.9
4-allylanisol - 25% -
Terpinen-4-ol 4.9 - 0.2
Tymol methyl ether 15 - 25.5
Dillapiol 1.5 25% 0.1
Garcia, 1999). In Folk veterinary, the aerial parts were
used as food integrator for rabbits and leaves as
galactogogue (Cornara et al., 2009). C. maritimum L.
leaves are rich in several compounds such as vitamin C,
caratenoids, flavonoids as well as bioactive substances
that could be used for aromatic, medicinal, antimicrobial
and insecticide. The oils extracted from leaves showed
the presence of high concentrations of fatty acids of the
ω-3 and ω-6 series. These fatty acids play an important
role in modulating human metabolism and have beneficial
effects against coronary heart diseases (Guil-Guerrero
and Rodriguez-Garcia, 1999). Recent works by Meot-
Duros et al. (2008) revealed that C. maritimum L. leaves
extract exhibited high phenol content and high ABTS
radical scavenging activity. The C. maritimum apolar
extract had strong antimicrobial activity against
Micrococcus luteus, Salmonella arizonae, Erwinia
carotovora, Pseudomonas fluorescens, P. aeruginosa, P.
marginalis, Bacillus cereus and Candida albicans (Meot-
Duros et al., 2008). Furthermore, C. maritimum L.
essential oil was studied, both for its antioxidant (Ozcan,
2000) and its antibacterial properties (Ruberto et al.,
2000). Essential oils had strong antibacterial action
against a large panel of human pathogenic bacteria
(Roosi et al., 2007). This is the case of the Gram-positive
bacteria B. cereus and M. luteus (Glowniak et al., 2006).
PHYTOCHEMISTRY
Volatile compounds
Both shoots and the fruit of C. maritimum L. are rich in
volatile compounds (Atia et al., 2009). The volatile oil
yield reaches about 0.8% in fruits and to 0.15 to 0.3% in
leaves (Franke, 1981). The major volatile oils identified
consist of sabinene, dillapiole, α-pinen, γ-terpinene
(crithmen), p-cymol, apiole, cis-ß-Ocimene, thymol and
terpinen-4-ol (Table 2). Other less abundant volatile
compounds were also found in C. maritimum L. such as:
α-Thujene, Camphene, α-Phellandrene, Limonene,
Cineole, trans-ß-Ocimene, trans-2-Ovten-1-ol,
Terpinolene, Linalool, trans-p-Menthen-1-ol, and
Myristicin. In leaves, the percentage of the major volatile
oils is highly variable: e.g. α-pinen (0.8 to 1.2%),
sabinene (33 to 40%), myrcene (1.6 to 1.8%), α-terpinene
(1.1 to 2.2%), p-cymene (3.7 to 9.3%), cis-ß-Ocimene (2
to 2.7%), γ -terpinene (22.3 to 28%), terpinen-4-ol (5 to
7.3%), the thymolmethylether (12.9 to 15.5%) and
dillapiole (1.1 to 3.1%) (Pateira et al., 1999). Concerning

3570 J. Med. Plant. Res.
the fruits, it has been reported that they contain
approximately 8 to 40% dillapiole, 12% α-pinen and up to
48% γ -terpinene (Franke, 1981).
Antioxidants, polyphenols and flavonoids
C. maritimum L leaves are rich in several compounds
such as vitamin C, caratenoids, and flavonoids (Guil-
Guerrero and Rodriguez-Garcia, 1999). Quantitative
analyses of the content of flavonoids, tannins and total
polyphenols in the aerial parts of C. maritimum L.,
showed that the content of flavonoids was 0.08 to 0.42%.
The tannin content ranged from 0.10 to 2.65%, while
the content of total polyphenols varied from 4.72 to
9.48%. The highest contents of flavonoids, tannins and
total polyphenols were found in the samples collected
before flowering and at the beginning of flowering (Males
et al., 2003). The fruit of C. maritimum L. was by high
accumulation of polyphenols. For instance, the endocarp
layer accumulated O-dihydroxyphenols (Atia et al., 2009
b).
The chlorogenic acid, a phenolic compound with high
radical-scavenging activity, was also found in
C.maritimum aerial parts (Meot-Duros and Magné, 2009)
Figure 3. The same authors identified the quinic acid,
another important phenolic compound. Quinic acid is a
cyclic polyol. It is produced synthetically by hydrolysis of
chlorogenic acid. This acid is used for the fabrication of
Tamiflu, a medicament for the treatment of influenza A
and B strains (http:/ /www.quinine-buchler. com/
quinicacid.htm). The aromatic ether O-geranylvanillin 3
was also isolated from C. maritimum L. (Cunsolu et al.,
1993). There is also evidence of some crithmic acid (p-
toluyl acid). In addition, polyacetylene compounds such
as falcarinon have been found in C. maritimum L.
(Hegnauer, 1973; Franke, 1981).
Recently, the occurrence of two bioactive flavonoids
was reported in C. maritimum L. Diosmin (3′, 5, 7-
trihydroxy-4′-methoxyflavone 7-rutinoside) and hesperidin
(3′, 5, 7-trihydroxy-4′-ethoxyflavanone7-rhamnoglycoside)
(Figure 2a and b) (Cornara et al., 2009). Meot-Duros et
al. (2010) purified the falcarindiol from the leaf apolar
extract of C. maritimum L. Falcarindiol is an antibacterial
and cytotoxic compound with multiple biological activities,
such as anti-inflammatory, antiplatelet-aggregatory and
antimutagenic properties (Miyazawa et al., 1996;
Christensen and Brandt, 2006). Falcarindiol extracted
from C. maritimum L. strongly inhibited the growth of M.
luteus and B. cereus. Moreover, this compound showed
cytotoxicity against IEC-6 cells (Meot-Duros et al., 2010).
Lipids
The oils extracted from C. maritimum L. leaves showed
the presence of high concentrations of fatty acids of the
ω-3 and ω-6 series (Guil-Guerrero and Rodriguez-
Garcia, 1999). On the dry weight basis, their percentage
reaches 2.02% for neutral lipids, 0.57% for the glycolipids
and 0.26% for the phospholipids. In the fruit, the
percentage of lipids reaches 44.4% on the dry weight
basis. C. maritimum L. fruit oil was also rich with oleic
acid (78.6%), low level of palmitic acid (4.8%) and non
negligible amount of linoleic acid (15.4%) (Atia et al.,
2010a). This composition is similar to olive oil and canola
oil. These results confirm the good quality of C.
maritimum L. oil.
Others
Several water-soluble compounds were observed in C.
maritimum. Among these solutes, the carbohydrates
namely sucrose and glucose, followed by organic acids
like malate and quinate (Meot-Duros and Magné, 2009).
C. maritimum L. leaves is a significant source of minerals;
hydrochlorates, sulphates, carbonates, potash, acetate,
iodine, and bromide (http://www.aromalves.com). Other
forms of minerals were also found in C. maritimum L. fruit
like phosphates, calcium, sulphur and sulphured amino
acids (Atia et al., 2010b).
CONCLUSION
In this review, we tried to present an updated overview
about the halophyte C. maritimum L. This plant has been
receiving the interest of the scientific community due to
its economical and fundamental interests: significant salt
tolerance in conjunction with potent medicinal and
economic importance. This species can grow in saline
land and irrigated with diluted sea water or diluted
brackish water. However, during germination and the
early growth stages, irrigating with non saline water is
needed. It is possible to successfully cultivate this
species in saline environments.
By studying the traditional uses and the
phytochemistry, we consider that C. maritimum L. is a
promising halophyte for biosaline agriculture and could
be proposed as a new industrial cash crop halophyte.
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