South African Journal of Botany 137 (2021) 216
227

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South African Journal of Botany

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Review

Nutritional and bioactive potential of seagrasses: A review

Doo Hwan Kima, Mohamad Fawzi Mahomoodallyb, Nabeelah Bibi Sadeerb, Park Gyun Seoka,
Gokhan Zenginc, Kishneth Palanivelood, Anees Ahmed Khalile, Abdur Rauff,
Kannan RR Rengasamyg,h,i,*
a
Department of Bioresources and Food Science, Konkuk University, Seoul 05029, Republic of Korea
b
Department of Health Sciences, Faculty of Medicine and Health Sciences, University of Mauritius, Re
duit, Mauritius
c
Department of Biology, Science Faculty, Selcuk University, Konya, Turkey
d
Institute of Ocean and Earth Sciences, Institute for Advanced Studies Building, University of Malaya,Wilayah Persekutuan Kuala Lumpur 50603, Malaysia
e
University Institute of Diet and Nutritional Sciences, Faculty of Allied Health Sciences, The University of Lahore, Pakistan
f
Department of Chemistry, University of Swabi, Swabi, Anbar, KPK, Pakistan
g
Institute of Research and Development, Duy Tan University, Da Nang 550000, Vietnam
h
Faculty of Environment and Chemical Engineering, Duy Tan University, Da Nang 550000, Vietnam
i
Indigenous Knowledge Systems Centre, Faculty of Natural and Agricultural Sciences, North-West University, Private Bag X2046, Mmabatho 2745, North West
Province, South Africa

A R T I C L E I N F O A B S T R A C T

Article History: Seagrasses, one of the most threatened yet overlooked ecosystems on Earth, are the only flowering plants to
Received 21 April 2020 recolonising the seabed. Apart from their critical ecological prominence on the life of many marine organ-
Revised 14 October 2020 isms, seagrasses are also used as an alternative or complementary medicine to manage an array of pathologi-
Accepted 19 October 2020
cal disorders such as muscle aches, wounds, abdominal pain, indigestion, hangover, and mental disorders.
Available online xxx
However, a compilation of existing work on their ethnopharmacological uses, nutritional values, pharmaco-
logical propensities and bioactive compounds is lacking. Thus, this review aims at elaborating on the bio-
Keywords:
chemical composition, phytochemical analysis, and biological properties including antioxidant, antimicrobial
Functional food
Halophytes
activities of various species of seagrasses. Seagrasses harbour several metabolites with multiple bioactivities.
Secondary metabolites The phytochemical compounds isolated from Zostera marina L., Thalassia testudinum K. D. Koenig. and Thalas-
Seagrasses sodendron ciliatum (Forssk.) exhibit a plethora of biological activities, including cytotoxicity against cancer
Bioactivities cell lines, anti-human immunodeficiency virus (HIV), antimicrobial, and skin regenerating properties. This
review also identifies vital lacuna in seagrass research. For instance, the mechanism and site of action of com-
pounds displaying potent biological activities has not been adequately addressed together with optimisation
of extraction methods to isolate minor metabolites and applying technological advancements in biological
assays. In conclusion, this review provides a synthesis of current knowledge and highlights future work that
needs to be undertaken for the biomedical application of such natural resources.
© 2020 SAAB. Published by Elsevier B.V. All rights reserved.

1. Introduction Kuttan, 2007) and neurodegenerative diseases such as Alzheimer’s

Since ancient times, traditional practitioners for treatment of vari-
ous diseases have utilized medicinal and herbal plants in primary
health care systems (Saleem et al., 2019a; Saleem et al., 2019b). Cur-
rently, scientists are focused on exploring phytochemicals present in
these medicinal plants as 80% of world’s population rely on these
conventional treatments for management of different diseases (Aziz
et al., 2014). The quest for bioactive compounds from plants has led
to the discovery of new pharmacophores and medicinal drugs which
offer efficient protection and treatment regimens against a plethora
of chronic pathologies, namely cancer (Kumar et al., 2004; Sheeja and
* Corresponding author.
E-mail address: rengasamyrrajakannan@duytan.edu.vn (K.R. Rengasamy).
(Mukherjee et al., 2007). To date, the health benefits of many phyto-
chemicals have been substantiated in the current pharmacological
record (Rouhi, 2003). For instance, pilocarpine derived from Pilocar-
pus jaborandi Holmes has been used as a clinical drug to treat glau-
coma for over 100 years (Dias et al., 2012), vancomycin isolated from
Amycolatopsis orientalis cultures is an approved Food and Drug
Administration (FDA) drug to treat severe infection (Jeong et al.,
2013).
Despite terrestrial plants proving their worth as a potential source
of bioactive compounds, the quest for novel prophylactic metabolites
should be an ongoing and continuous task, since new diseases keep
emerging and existing diseases keep changing their mechanisms of
action (Malve, 2016). Alternative sources of plants with different

https://doi.org/10.1016/j.sajb.2020.10.018
0254-6299/© 2020 SAAB. Published by Elsevier B.V. All rights reserved.
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al.
phytochemical properties should be considered. Research in marine
pharmacology is still scanty although the marine environment
houses a diverse array of organisms which could serve as a potential
source of pharmacologically active compounds (Alice and Ele-
gbede, 2016; Malve, 2016).
Marine compounds are trivial in the current pharmacopoeia;
however, it is anticipated that the marine environment could offer a
new scope of research on marine-derived drugs as it represents 95%
of the biosphere (Danovaro et al., 2017; Jimeno et al., 2004). Plants
from the aquatic ecosystem have not been fully explored due to the
complexity of the marine ecosystem with inter-connected food
chains, constant change in environment and variation in marine spe-
cies. One such group of underexplored marine plants are the sea-
grasses (Rengasamy et al., 2019; Ziegler, 2010).
Seagrasses, categorized as halophytes, are rooted, marine angio-
sperms capable of completing their life cycle whether permanently
or partially submerged in water (Aboud and Kannah, 2017). They are
the only flowering plants to recolonise the seabed and, play a signifi-
cant role in supporting food security and mitigating climate change
(Unsworth et al., 2019). Seagrasses form important ecosystems for
millions of marine organisms and help in the stabilisation of sedi-
ments (Larkum et al., 2006). They act as an important feeding and
breeding ground for several threatened organisms, including
dugongs, turtles and sea horses. Seagrass beds are inhabited by a
range of benthic organisms such as sea cucumbers, sea urchins, star-
fishes, clams and sponges (Wahab et al., 2017). The global seagrass
diversity is recorded at 72 species (Duffy et al., 2019). Relatively low
levels of direct grazing have prompted the studies on the chemical
constituents and relative food value of seagrasses (Dawes and Law-
rence, 1983). Especially coastal populations (Heminga and Durate,
2000) use seagrass biomass as human food. It can be a rich source of
copper when consumed directly (de Los Santos et al., 2019). In folk
medicine, seagrasses have been used for a variety of remedial pur-
poses, for example in the treatment of fever, skin diseases, muscle
pains, wounds, stomach problems, as tranquillizers for babies as a
remedy against ray stings (de la Torre-Castro and Ro € nnba
€ck, 2004)
and as a potential antimicrobial agent (Kannan et al., 2013). Halophila
ovalis was used by the fishing communities of Cuddalore and Naga-
pattinam districts of Tamil Nadu, South India as medicine to treat var-
ious skin diseases, burns and boils (L Kannan et al., 1999). It is also
used as a potent antioxidant and anti-inflammatory agent (Yuvaraj
and Arul, 2018). The seeds of the tropical seagrass Enhalus acoroides
were traditionally eaten in the Philippines (Klangprapun et al., 2018;
Montan ~ o et al., 1999) and were thought to have aphrodisiac and con-
traceptive properties (Alino et al., 1990). Available scientific data con-
firms an array of promising biological activities of E. acoroides,
namely antimicrobial, insecticidal, antimalarial, vasoprotective prop-
erties, anti-inflammatory, antioxidant and antialgal (De Vincenti
et al., 2018). Nonetheless, as far as our literature search could reach,
no review has been compiled yet elaborating on the promising bio-
medical framework exhibited by seagrasses. Therefore, this present
review amalgamates a comprehensive compilation of the phyto-
chemical composition and biomedical applications of seagrasses
around the world.
2. Biochemical composition
The seeds of E. acoroides were analysed for their nutritive value to
assess their dietary value for human. The raw flour prepared from
dried seeds was subjected to different an array of analysis to deter-
mine its nutritional content. They contained 9.8% moisture, 8.8% pro-
tein, 0.2% fat, 72.4% carbohydrate, 2.4% crude fibre, 6.4% ash, calcium
(933 mg kg
1), phosphorus (2392 mg kg
1) and iron (2813 mg kg
1).
These values were similar to plants of terrestrial origin
(Montan ~ o et al., 1999); (Geneid and El-Hady, 2006). The crude pro-
tein and ash contents in Ruppia cirrosa were 11.94 and 16.78%,
217
South African Journal of Botany 137 (2021) 216
227
respectively and in Z. marina, the contents were 11.31 and 19.20%,
respectively. The hydrolysed polysaccharides detected in both sea-
grass species were galactose, glucose, arabinose, xylose, rhamnose
and uronic acid with glucose being the most abundant (Milchakoya
et al., 2014). Seasonal level of ash, protein, carbohydrate and calorific
values of blades belonging to three species of seagrasses Thalassia tes-
tudinum, Syringodium filiforme and Halodule wrightii collected at
seven sites on the west coast of Florida were investigated
(Dawes, 1986); (Dawes and Lawrence, 1983, 1980). The macro (phos-
phorus, potassium, calcium, magnesium, sodium) and micro (nickel,
zinc, iron, chromium, copper, cadmium, lead) nutrients, protein, car-
bohydrate, lipid, chlorophyll, ash and fibre compositions from three
seagrasses namely Cymodocea serrulata, Halodule uninervis and
Syringodium isoetifolium collected from the Gulf of Mannar were
compared. The lipid and ash contents were higher in H. uninervis
while S. isoetifolium exhibited lower composition level. However,
micronutrients and calcium were higher in S. isoetifolium (690 mg/
100 g) in contrast to the other two seagrasses (Jeyasanta et al., 2018).
The proximate compositions of protein, fat, ash, crude fibre, car-
bohydrate and calories in turtle grass (Thalassia testudinum) was sim-
ilar to Coastal Bermuda grass (Burkholder et al., 1959; (Drew, 1983).
Significant quantitative differences were noticed amongst sterols
(sitosterol and stigmasterol) and terpenoids (campesterol and choles-
terol) contents of the seagrass Porteresia coarctata that were grown in
two different mangrove habitats of Sundarbans (Misra et al., 1987).
High salinity significantly decreased the protein content in P. coarc-
tata (Ghosh and Mitra, 2015). Salinity is a major factor that dominates
the growth, structure, function and distribution of seagrasses in the
oceans (Oscar et al., 2018). Cymodocea rotundata was rich in nitrogen
(6.19%), organic carbon (30.8%) and major elements compared to
marine algae. The average sodium (Na), potassium (K), and magne-
sium (Mg) composition of this seagrass species were relatively higher
(7758, 842.2 and 1412.6 mg/g, respectively) compared to the Ca con-
tent (46.8 mg/g) (Jagtap and Untawalw, 1984). This was consistent
with previous studies where seagrasses are usually rich in Na, K and
Mg (Jagtap and Untawale, 1984). The major structural constituents of
the seagrasses Posidonia australis cell wall are cellulose and lignin
(Torbatinejad et al., 2007). Athiperumalsami et al. (2008) analysed
phytochemical constituents of six genera of seagrasses from the Gulf
of Mannar contained medicinally important primary (sugar, amino
acids, and fats & oils) and secondary (catechin, alkaloids, coumarin,
flavonoids, steroid, tannin, and glycosides) bioactive compounds
(Athiperumalsami et al., 2008). The calorific value of the Indian sea-
grasses was equivalent to Bengal gram, peas, potatoes and sweet
potato (Pradeepa et al. 2011). The detailed study on the nutritional
composition of seagrasses of India showed that seagrasses of this
region have enormous potential for the development of functional
food ingredients and pharmaceutical supplements (Kannan et al.,
2013). Compilation on the biochemical composition of seagrasses are
shown in Table 1.
3. Phytochemical analysis
Many studies on the occurrence of specific chemical compounds
in seagrasses have been undertaken. In most cases, studies have been
directed towards seagrass phylogeny (Gillan et al., 1984). For exam-
ples, the distribution of phenolic acids, sulphated phenolic com-
pounds and flavones sulphates in seagrasses show marked family
and the generic differences (McMillan et al., 1980). As part of a pro-
gramme directed towards the elucidation of carbon flow in food
chains in temperate seagrass communities, lipids were used as dis-
tinctive chemical markers for the seagrasses Posidonia australis and
Heterozostera tasmanica. The most abundant lipid classes were fatty
acids and there were minor constituents in H. tasmanica. (Nichols
et al., 1982). Phytochemical profiling of E. acaroides methanolic
extract by liquid chromatography/mass spectrometry (LC/MS)
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al. South African Journal of Botany 137 (2021) 216
227

Table 1
Comparison of means or ranges of concentrations of chemical composition of seagrassses.

Carbohydrate Protein Lipid Fibre Ash Phenol Flavonoid Tannin References

2.018
7.258a 5.61
14.404a 1.40
9.56a 16.5
25.2a 18.4
27.366a 0.395
0.655b 0.112
0.73b 0.748
9.731b Kannan et al., 2013
2.0
8.7b 0.1
5.9b 0.01
3.2a

3.2
9.2b
0.56
2.4b Pradheepa et al., 2011
6.015
6.327a 6.796
7.893a 0.92
1.02a




Kannan et al., 2010
0.140
3.180b 2.63
15.75b 0.022
0.030b 15.1
57.3b
0.070
0.226b
2.94
9.8b Athiperumalsamy et al., 2008




151
200c

17.4
18.5c Torbatinejad et al., 2007


0.17
0.71a 13.5
26.7a 13.8
42.9a


Yamamuro et al., 2005
72.4
87.6 a 9.8
11 a 0.1
0.2 a 0.4
2.4 a 0.5
6.4 a


Montan ~ o et al., 1999

10.4
11.5 a 0.5
1.5 a
23
44 a


Dawes et al., 1987
65.3 a 11.3 a 1.4 a 7.4 a 7.1 a


Valencia et l., 1985
11.1
20a 7
10a 1.2
3.8a 34
41a Dawes., 1986

5.57
12.50a

4.0
5.9a


Augier et al., 1982
6
50a 5
22a 0.5
4.0a
25
56a


Dawes and Lawrence, 1980

10.0a 0.8a




Park, 1969
22.95
33.068a 1.03
2.61a 3.5
5.06a 0.144
0.835b 0.077
0.501b 0.225
1.347b 0.736
2.45b Kannan et al., 2013



0.02
0.19 0.03
0.149 0.05
0.29
Pradheepa et al., 2011



0.454
0.885b 0.247
0.541b 0.701
1.424b 0.508
2.39b Kannan et al., 2010
1.76
3.35a 5.6
22.28b
0.385
1.043b 0.161
0.719b
0.093
0.30b Athiperumalsamy et al., 2008
17.7
38.9a 0.41
3.02a




Yamamuro et al., 2005

0.95
2.1a




Terrados et al., 1999a

1.84
2.02




Terrados et al., 1999b
33.6a 1.92a




Duarte, 1990
31.45
36.85 a 1.30
2.75 a 4.40
5.12 a



Augier et al., 1982
a
%;.
b
mg g
1;.
c
g kg
1.

detected the flavones apigenin and luteolin, three kaempferol deriva-
tives and azelaic acid (De Vincenti et al., 2018). Detailed examinations
of specific lipids in Zostera muelleri, using high-performance capillary
gas chromatography and mass spectroscopy found that out of the 30
sterols identified from the sediments, 24-methylcholesta-5.22E‑dien-
3b-ol was the major steroid compound (Volkman and Johns, 1977).
Detailed analysis of other seagrasses using capillary gas chromatogra-
phy/mass spectrometry (GC/MS) is lacking. Packed column GC analy-
sis of fatty acids and sterols in a few seagrasses showed b-sitosterol
as the major constituent in Halophila engelmanni, Thalassia testudi-
num and Ruppia maritima and stigmasterol predominating in Diplan-
thera wrightii and Syringodium filiforme (Maurer and Parker, 1967;
Attaway et al. 1971).
Phenolic compounds are widely distributed in marine plants,
and several biological functions have been attributed to them
(Harborne and Williams, 1976; McMillan et al., 1980; Zapata and
McMillan, 1979). Examples of the natural products isolated from
Zostera marina L. (eelgrass) include flavone sulphates
(Harborne and Williams, 1976), the 7-sulphates of luteolin, dio-
smetin, apigenin, chrysoeriol, the 7,30 - disulphate of luteolin and
several non-sulphated phenolic acids; p-coumaric, ferulic, caffeic,
vanillic, gallic, protocatechuic, gentisic,4‑hydroxy benzoic and o-
pyrocatechuic acids (Quackenbush et al., 1986; Zapata and McMil-
lan, 1979). A total of six phenolic acids were quantified in the
water-soluble extracts of Zostera marina L. from Puget Sound,
Washington using high-performance liquid chromatography
(HPLC) techniques (Quackenbush et al., 1986). Halophila ovalis of
the Pacific Ocean contains sulphated flavonoids (McMillan, 1986).
Leaf extracts of eelgrass containing phenolic constituents may
inhibit amphipod grazing, microbial growth (Harrison, 1982) and
photosynthetic carbon uptake in epiphytic diatoms (Harrison and
Durance, 1985). It has been suggested that phenolic acids may
confer resistance to the so-called 'wasting disease' that was
blamed for the catastrophic die-off of eelgrass in the North Atlan-
tic in the 1930s (Buchsbaum et al., 1990). The phytochemical
composition of the dried powdered H. ovalis was analysed quanti-
tatively using GC/MS for alkaloids, carbohydrates, flavonoids, phe-
nols, saponins, and tannins. Phenols (9.44 § 0.02 mg/g) were
most abundant followed by saponins (9.07 § 0.06 mg/g), flavo-
noids (5.59 § 0.01 mg/g), proteins (5.19 § 0.02 mg/g),
carbohydrates (4.82 § 0.04 mg/g), alkaloids (1.06 § 0.03
mg/g) and tannins (0.26 § 0.02 mg/g) (Yuvaraj et al., 2012).
Simple phenolic acids are abundant and account for as much as
half of the measured total phenolics in seagrasses (Vergeer et al.,
1995; Vergeer and Develi, 1997). For example, H. ovalis yielded phe-
nols in significant amounts compared to other phytochemicals quan-
tified (Yuvaraj et al., 2012). The array of phenolic acids produced by
seagrasses is dependant upon the species, population, and tissue
examined. Caffeic acid, protocatechuic acid, p-coumaric acid, p-
hydroxybenzoic acid, ferulic acid, vanillic acid, and gentistic acid are
amongst others reported (McMillan et al., 1980; Quackenbush et al.,
1986; Ravn et al., 1994). Phenolic acid conjugates are also common in
seagrasses. For instance, sulphated conjugates were present in all 43
species examined (McMillian et al., 1980). Concentrations of phenolic
acids vary dramatically and are closely connected to both biotic and
abiotic factors. For example, concentrations of rosmarinic acid and
caffeic acid in Zostera marina, varied seasonally, ranging from 0.4 to
19.2 mg/g dry mass (DM) (Harrison and Durance, 1989; Ravn et al.,
1994). A comparative study showed that the rhizomes of Posidonia
oceanica yielded a higher level of phenols in summer in contrast to
winter (Baby et al., 2017).
Seagrasses also accumulate significant levels of condensed tan-
nins (i.e., proanthocyanidins) in leaf and shoot tissues. For example,
3
11% have been recorded in the leaves of the turtlegrass, Thallasia
testudinum, from the Gulf of Mexico and Caribbean Sea (Arnold and
Targett, 2002), and proanthocyanin levels vary significantly over brief
periods (Arnold and Targett, 2002). However, mesograzers such as
Crepidula ustulatulina and Paracerceis caudata may affect the compo-
sition of phenolic acids and proanthocyanidins of various seagrasses
(Steele and Valentine, 2015).
A comparison between secondary compounds of leaves of Hetero-
zostera tasmanica collected from Australia and Chile found similar
phenolic compound profiles in samples from both sites (McMil-
lan, 1983). The 43 species of seagrasses analysed consistently con-
tained either flavone, phenolic acid sulphates or a combination of
both. Five of the twelve genera analysed had sulphated flavones
while Zostera and Enhalus lacked the sulphated phenolic acids
(McMillan et al., 1980). Growth of Z. marina was monitored in run-
ning seawater mesocosms with either a mud or sand substratum and
the phenolic-nitrogen interactions investigated. The leaf nitrogen

218
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al.
and phenolic contents were inversely related. Seagrass with a natu-
rally low phenolic content were prone to total mortality from wast-
ing disease caused by a slime-mould-like parasite if grown in an
environment with a high nitrogen composition e.g. mud. On the con-
trary, a high phenolic content prevented mortality (Buchsbaum et al.,
1990).
p-(sulphooxy) cinnamic acid was isolated for the first time from Z.
marina . It was beneficial for the seagrass since the compound pre-
vents attachment of marine bacteria and barnacles to artificial
surfaces at nontoxic concentrations. Volatile compounds (octadeca-
namide, nonadecane, hexadecanamide, phytol, heptadecane, and
pentadecane) were identified in the oil from the shoots of Z. marina
using GC and GC/MS (Kawasaki et al., 1998). The fatty acids myristic
acid, pentadecanoic acid, palmitic, margaric acid, stearic acid, arachi-
dic acid, behenic acid, lignoceric acid, palmitoleic acid, oleic acid, hex-
adecatrienoic acid, arachidonic acid, eicosapentaenoic acid,
docosahexaenoic acid, linoleic acid and a-linolenic acid were isolated
from Z. marina (Custodio et al., 2016). LC/MS analysis detected p-cou-
maric acid, caffeic acid, ferulic acid, zosteric acid, three sulphated fla-
vonoids namely apigenin-7-sulfate, luteolin-7-sulfate and diosmetin-
7-sulfate from the same seagrass species (Papazian et al., 2019).
Recently, cymodienol, a cyclic diarylheptanoid was isolated from Z.
marina, which had only previously been detected from the seagrass
Cymodocea nodosa (Li et al., 2019). However, similar to other sea-
grasses, the phenolic profiles of Z. marina vary with geographic
regions (Grignon-Dubois and Rezzonico, 2012).
Two diarylheptanoids, cymodienol and cymodiene were identi-
fied in Cymodocea nodosa collected off the coastal areas of central
Greece as the first members of this class of compounds isolated from
marine organisms (Kontiza et al., 2005). Secondary metabolites have
been studied in seagrasses for chemotaxonomic purposes
(Larkum et al., 2006). Phytochemical screening revealed the presence
of two novel prenylated flavon-di-O-glycosides namely cymodioside
A and B from C. nodosa (Smadi et al., 2018). An investigation into the
phenolic constituent of C. nodosa revealed the presence of seven
identified phenolics including sinapinic acid derivative, catechin,
ferulic acid derivative, cinnamic acid derivative, quercetin-3-O-ruti-
noside, quercetin-3-O-glucoside and isorhamnetin-3-O-rutinoside
(Kolsi et al., 2017). Chicoric acid was identified in the mature leaves
of Posidonia oceanica, in high concentrations together with p-cou-
maric acid, vanillin, ferulic acid, and gentisic acid (Haznedaroglu
and Zeybek 2007). Large amounts of catechins were detected in
P. oceanica (Leri et al., 2018). In two different studies, the inosi-
tols, a class of cyclitol from the flotsam from the seagrass Syringo-
dium filiforme (Nuissier et al., 2008) and glucosyl terpenoid
macrocycle, syphonoside from Halophila stipulacea (Carbone et al.,
2008) were characterized.
The response to wounding induces accumulation of tannin in tur-
tle grass, Thalassia testudinum (Arnold et al., 2008). In an experimen-
tal design to evaluate the effects of wounding on tannin
accumulation, the turtle grass tissue wounded by fish grazing, graz-
ing by the urchin Lytechinus variegatus, and treatment with the natu-
ral plant wound hormone jasmonic acid was analysed. Fish grazing
triggered the accumulation of 10 mg tannin g
1 dry mass (DM) after
5 days. Tannin accumulation correlated with a reduction of blade
extension rates (Arnold et al., 2008). Zosteric acid (ZA) (Achamlale
et al., 2009a) and rosamarinic acid (Achamlale et al., 2009b) was iso-
lated from the detritus of Zostera noltii and Zostera marina collected
from the beaches of Arcachon lagoon (France). Chemical analysis
of Halophila stipulacea yielded a malonylated flavone glucoside,
genkwanin-40 -O-(6“-malonyl-glucopyranoside) along with six fla-
vone glucosides derivatives which were previously only reported
from terrestrial source. This report highlighted the first for malo-
nylated metabolites in marine phanerogams (Bitam et al., 2010).
Table 2 provides a list of all compounds reported from seagrasses
to date.
219
South African Journal of Botany 137 (2021) 216
227
4. Biological properties
Trends in secondary metabolites research are often leading
towards drug discovery with an emphasis on bioactivity. Exploration
of the marine ecosystem produces numerous complex and novel
chemical entities. Due to the physically harsh condition in the marine
environment, marine organisms are adapted to produce a variety of
molecules with unique structural features, which exhibit various bio-
logical activities (Ravikumar et al., 2010). Seagrasses are well docu-
mented for the presence of diverse potent secondary metabolites
(Puglisi et al., 2007) with documented bioactivity such as antimicro-
bial, antioxidative and antifouling. There is a scarcity of studies avail-
able on antifungal, antibacterial and antiviral activities of crude
solvent extracts of seagrasses. Most of the works carried out so far
are all in combination with seaweeds and other plants. The following
section complies seagrass related bioactivity. Table 3 compiles a sum-
mary of seagrass related bioactivity.
4.1. Antimicrobial properties
Extracts of some seaweeds, seagrasses and lemongrass oil from
the Papua New Guinea were tested for their antimicrobial activities.
The methanol and hexane extracts of Enhalus acoroides and some sea-
weeds were tested against Bacillus subtilis, Staphylococcus aureus,
Pseudomonas aeruginosa and Klebsiella pneumoniae. The seagrass
methanolic extract inhibited the growth of all tested bacteria except
Klebsiella pneumoniae (Alam et al., 1994). Halophila ovalis, Halophila
beccarii and Halodule pinifolia from Porto Novo were tested against
bacterial pathogens. Halodule pinifolia exhibited more promising
activity against all tested pathogens (Balasubramanian et al., 2000).
The antifouling potential of some marine organisms against Bacillus
and Pseudomonas species showed that Cymodocea rotundata exhib-
ited mild activity against all the bacterial strains (Bhosale et al.,
2002).
Four new metabolites (deoxycymodienol, isocymodiene, nodosol,
and briarane diterpenes) were isolated from the organic extract of
Cymodocea nodosa collected and evaluated for their antibacterial
activity against multi-drug resistant pathogens including methicillin-
resistant strains of Staphylococcus aureus and the rapidly growing
mycobacterium (Mycobacterium phlei, M. smegmatis and M. fortui-
tum) (Kontiza et al., 2008). amongst these four new isolated metabo-
lites, nodosol showed best results in terms of experimented
antimicrobial activity assay. The geographical effects of antimicrobial
property in the extracts of two seagrass species, namely E. acoroides
and Halophila minor against marine pathogens and saprophytes were
evaluated. amongst these examined seagrass species, E. acorioides
extracts more showed significantly more antimicrobial activity
against P. bacteriolytica, S. aggregatum, and D. salina as compared to
H. minor (Puglisi et al., 2007). Antibacterial properties of three sea-
grasses, namely Cymodacea serrulata, Halophila ovalis and Zostera
capensis were tested against human pathogens Staphylococcus aureus,
Bacillus cereus, Bacillus subtilis, Escherichia coli, Salmonella paratyphi,
Salmonella typhimurium and Micrococcus luteus. The ethyl acetate and
methanolic extracts exhibited the highest antibacterial activities
(Kumar et al., 2008). The chemical constituents of the ethanol
extracts of E. acoroides were characterized and evaluated for antifee-
dant, antibacterial and antilarval activities. Eleven pure compounds
including four flavonoids and five sterols were detected which dis-
played potent antifeedant, antibacterial and antilarval activities (Qi
et al., 2008). As antibacterial biarane diterpene was isolated from
Cymodocea nodosa (Kontiza et al., 2008). The methanol extracts of
Syringodium isoetifolium were screened against bacterial and fungal
pathogens which cause disease to silkworm Bombyx mori and maxi-
mum inhibition were recorded against Proteus vulgaris (Ravikumar
et al., 2010). Comparison of the antimicrobial potential of three sea-
grasses, namely Halophila stipulacea, Halodule pinifolia and
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al. South African Journal of Botany 137 (2021) 216
227

Table 2
Compounds found in seagrasses.

Compounds Seagrasses Biological activity References

D-Allose (1) H. pinifolia
Ragupathi Raja Kanna et al., 2012

4H-Pyran-4-one-2,3-dihydro-3,5-dihy- H. pinifolia
Ragupathi Raja Kanna et al., 2012
droxy-6-methyl-
5-Caranol, trans,trans-(+)- H. pinifolia
Ragupathi Raja Kanna et al., 2012
Galactinol dihydrate luteolin H. pinifolia
De Vincenti et al., 2018
Procyanidin B2, H. pinifolia
De Vincenti et al., 2018
+(
) epicatechin H. pinifolia
De Vincenti et al., 2018
Quercetin-3-(6-malonyl)-glucoside H. pinifolia
De Vincenti et al., 2018
Quercetin H. pinifolia
De Vincenti et al., 2018
Azelaic acid H. pinifolia
De Vincenti et al., 2018
Apigenin H. pinifolia
De Vincenti et al., 2018
Isorhamnetin H. pinifolia
De Vincenti et al., 2018
Rutin T. ciliatum Antioxidant, Antiviral and Cytotoxicity Hamdy et al., 2012
against cancer cell lines
Asebotin, T. ciliatum Antioxidant, Antiviral and Cytotoxicity Hamdy et al., 2012
against cancer cell lines
3-hydroxyasebotin T. ciliatum Antioxidant, Antiviral and Cytotoxicity Hamdy et al., 2012
against cancer cell lines
Quercetin-3-O-ß-D-xylopyranoside T. ciliatum Antioxidant, Antiviral and Cytotoxicity Hamdy et al., 2012
against cancer cell lines
Catechin T. ciliatum Antioxidant, Antiviral and Cytotoxicity Hamdy et al., 2012
against cancer cell lines
Phenol P. oceanica
Cuny et al., 1995; Agostini et al., 1998
Phloroglucinol P. oceanica
Cuny et al., 1995; Agostini et al., 1998
Pyrocatechol P. oceanica
Cuny et al., 1995; Agostini et al., 1998
Pyrogallol P. oceanica
Cuny et al., 1995; Agostini et al., 1998
4-Hydroxybenzaldehyde P. oceanica
Cuny et al., 1995; Agostini et al., 1998
3,4-Dihydroxybenzaldehyde P. oceanica
Cuny et al., 1995; Agostini et al., 1998
Syphonosides 1 H. stipulacea
Gavagnin et al., 2007; Bitam et al., 2010
Syphonosides 2 H. stipulacea
Bitam et al., 2010
Genkwanin-40 -O-ß-(60 ’-O-malonyl- H. stipulacea
Bitam et al., 2010
glucopyranoside)
Enkwanin-40 -O-ß-glucopyranoside H. stipulacea
Bitam et al., 2010
Apigenin E. acoroides Antifeedant, antibacterial, and antilarval Qi et al., 2008
activities
Apigenin E. acoroides
De Vincenti et al., 2018
Luteolin E. acoroides
De Vincenti et al., 2018
Azelaic acid E. acoroides
De Vincenti et al., 2018
Apigenin-7-ß-(60 ’-O-malonyl- H. stipulacea
Bitam et al., 2010
glucopyranoside)
Apigenin- 40 -O-ß-glucopyranoside H. stipulacea Bitam et al., 2010
Chrysoeriol-40 -O-ß-glucopyranoside H. stipulacea Bitam et al., 2010
Glucopyranoside H. stipulacea Bitam et al., 2010
Chrysoeriol-7-O-ß-glucopyranoside H. stipulacea Bitam et al., 2010
Vanillin P. oceanica Cuny et al., 1995; Agostini et al., 1998;
Haznedaroglu and Zeybek, 2007
Syringa aldehyde P. oceanica Cuny et al., 1995; Agostini et al., 1998
Syringin T. hemprichii Qi et al., 2012
Benzoic acid P.oceanica Cuny et al., 1995; Agostini et al., 1998
p-Hydroxybenzoic acid P. oceanica, C. nodosa, Z. marina, T. testu- Cariello et al., 1979; Cuny et al., 1995;
dinum, T. hemprichii Agostini et al., 1998; Dumay et al.,
2004; Nuissier et al., 2010; Zapata &
Mac Millan, 1979; Quackenbush et al.,
1986; Bushmann and Ailstock (2006);
Qi et al., 2012
p-Anisic acid P. oceanica Cuny et al., 1995; Agostini et al., 1998
Protocatechuic acid P. oceanica, T. testudinum Cuny et al., 1995; Agostini et al., 1998;
Zapata & Mac Millan, 1979
Vanillic acid P. oceanica, Z. marina, T. testudinum Cariello et al., 1979; Cuny et al., 1995;
Agostini et al., 1998; Nuissier et al.,
2010; Zapata & Mac Millan, 1979
4-O-p-Hyxdroxybenzoyl-vanillic acid P. oceanica Cariello et al., 1979
glucopyranosyl ester
Gallic acid P.oceanica, Z. marina Agostini et al., 1998; Quackenbush et al.,
1986
Syringic acid P. oceanica Cuny et al., 1995; Agostini et al., 1998;
Nuissier et al., 2010; Zapata & Mac
Millan, 1979
Gentisic acid P. oceanica, Z. marina Haznedaroglu and Zeybek, 2007;
Quackenbush et al., 1986
Zosterin Z. asiatica, Z. marina Loenko et al., 1997; Dolgii et al., 1999;
Khasina et al., 2003
p-coumaric acid Z. marina
Papazian et al. (2019)
Caffeic acid Z. marina

(continued)

220
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al. South African Journal of Botany 137 (2021) 216
227

Table 2 (Continued)

Compounds Seagrasses Biological activity References

Ferulic acid Z. marina

Zosteric acid Z. marina

Apigenin-7-sulfate Z. marina

Luteolin-7-sulfate Z. marina

Diosmetin-7-sulfate Z. marina

Cymodienol Z. marina
Li et al., (2019)
Cymodienol C. nodosa Antibacterial activity against methicillin- Kontiza et al., 2005 & 2008
resistant (MRSA) strains
Cymodiene C. nodosa Kontiza et al., 2005
3-keto steroids C. nodosa Kontiza et al., 2006
Cymodioside A and B C. nodosa
Smadi et al., 2018
Snapinic acid derivative C. nodosa
Kolsi et al., 2017
Catechin C. nodosa

Ferrilic acid derivative C. nodosa

Cinnamic acid derivative C. nodosa

Quercetin-3-O-rutinoside C. nodosa

Quercetin-3-O-glucoside C. nodosa

Isorhamnetin-3-O-rutinoside C. nodosa

Subfraction H5 Z. japonica Hua et al., 2006
Thalassiolins A (luteolin 7-O-ß-D-gluco- T. testudinum, T. hemprichii Antifouling activity, Anti-HIV activity Jensen et al., 1998; Rowley et al., 2002; Qi
pyranosyl-20 -sulfate) et al., 2012
Thalassiolins B T. testudinum, T. hemprichii Anti-HIV activity, antioxidant and skin- Rowley et al., 2002; Regalado et al., 2009;
regenerating activities Qi et al., 2012
Thalassiolins C T. testudinum Anti-HIV activity Rowley et al., 2002
Apigenin-7
0-b-D-glucoside, Z. marina, H. stupulaceae Kim et al., 2004; Bitam et al., 2010
Chrysoeriol Z. marina Kim et al., 2004
Luteolin Z. marina, E. acoroides Kim et al., 2004; Qi et al., 2008
Luteolin 40 -glucuronide
Luteolin 30 -glucuronide
Thalassiolins A (luteolin 7-O-ß-D-gluco- T. testudinum Antifouling activity Jensen et al., 1998
pyranosyl-20 -sulfate)
L-chiro-inositol S. filliforme Nuissier et al., 2008
Zosteric acid Z. noltii Achamlale et al., 2009a
Rosmarinic acid Z. marina Achamlale et al., 2009b
Caftaric acid Nuissier et al., 2010
p-OH-Benzoic Nuissier et al., 2010
Coutaric Nuissier et al., 2010
Di-coumaroyl tartaricc Nuissier et al., 2010
Di-feruloyl-tartaric Nuissier et al., 2010
Piceol P. oceanica Agostini et al., 1998
Acetovanillone P. oceanica Agostini et al., 1998
Acetosyringone P. oceanica Cuny et al., 1995; Agostini et al., 1998
Catechin P. oceanica
Leri et al., 2018
Cinnamic acid P. oceanica; Antifouling activity Cuny et al., 1995; Agostini et al., 1998;
Haznedaroglu and Zeybek (2007);
Todd et al. 1993; Dumay et al. (2004);
Bushmann and Ailstock (2006)
Coumaric acid P. oceanica, Z. marina; T. testudinum, H. Antioxidant activity Cuny et al., 1995; Agostini et al., 1998;
pinifolia, T. hemprichii Haznedaroglu and Zeybek (2007);
Dumay et al. (2004); Achamlale et al.,
2009a; Bushmann and Ailstock (2006);
Zapata & Mac Millan, 1979; Ragupathi
Raja Kannan et al., 2010
Ferulic acid P. oceanica, Z. marina, T. testudinum Cuny et al., 1995; Agostini et al., 1998;
Haznedaroglu and Zeybek (2007);
Bushmann and Ailstock (2006); Dumay
et al. (2004); Quackenbush et al., 1986;
Zapata & Mac Millan, 1979
Sinapic acid P. oceanica Cuny et al., 1995; Agostini et al., 1998;
Haznedaroglu and Zeybek (2007);
Dumay et al. (2004);
Quackenbush et al., 1986
Caffeic acid P. oceanica, Z. marina, T. testudinum, T. Antioxidant, Antiviral and Cytotoxicity Cuny et al., 1995; Agostini et al., 1998;
hemprichii, T. ciliatum against cancer cell lines Haznedaroglu and Zeybek (2007);
Bushmann and Ailstock (2006); Dumay
et al. (2004); Nuissier et al., 2010;
Zapata & Mac Millan, 1979; Qi et al.,
2012; Hamdy et al., 2012
Chlorogenic acid P. oceanica, C. nodosa Cariello et al., 1979)
Cichoric acid P. oceanica, S. filiforme, T. hemprichii Haznedaroglu and Zeybek (2007); Car-
iello and Zanetti (1979); Nuissier et al.,
2010; Qi et al., 2012
Phloretin P. oceanica Cuny et al., 1995
Phloridzin P. oceanica Cuny et al., 1995
Kaempferol P. oceanica Cannac et al. (2006, 2007).

(continued)

221
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al. South African Journal of Botany 137 (2021) 216
227

Table 2 (Continued)

Compounds Seagrasses Biological activity References

Quercetin P. oceanica Cannac et al. (2006, 2007).

Isorhamnetin P. oceanica Cannac et al. (2006, 2007).
Myricetin P. oceanica Cannac et al. (2006, 2007).
5a-Cholestan-3-ol P. oceanica, C. nodosa Sica et al. (1984)
5a-Cholest-22E-en-3-ol P. oceanica, C. nodosa Sica et al. (1984)
5a-Campestanol P. oceanica, C. nodosa Sica et al. (1984)
5a-Ergostanol P. oceanica, C. nodosa Sica et al. (1984)
5a
24-Methylcholest-24(28)-en-3-ol P. oceanica, C. nodosa Sica et al. (1984)
5a-b-Sitostanol P. oceanica, C. nodosa Sica et al. (1984)
5a-Isofucostanol P. oceanica, C. nodosa Sica et al. (1984)
Cholesterol P. oceanica, C. nodosa Sica et al. (1984)
Desmosterol P. oceanica, C. nodosa Sica et al. (1984)
Cholesta-5,22-diene-3b-ol P. oceanica, C. nodosa Sica et al. (1984)
Campesterol P. oceanica, C. nodosa Sica et al. (1984)
Dihydrobrassicasterol P. oceanica, C. nodosa Sica et al. (1984)
Epibrassicasterol P. oceanica, C. nodosa Sica et al. (1984)
Brassicasterol P. oceanica, C. nodosa Sica et al. (1984)
24-Methylenecholesterol P. oceanica, C. nodosa Sica et al. (1984)
b-Sitosterol P. oceanica, C. nodosa Sica et al. (1984)
Isofucosterol P. oceanica, C. nodosa Sica et al. (1984)
Stigmasterol P. oceanica, C. nodosa Sica et al. (1984)
Stigmasta-4,22‑dien-6b-ol-3-one E. acoroides antifeedant, Qi et al., 2008
Stigmasta-4,22‑dien-3,6-dione E. acoroides antibacterial, and antilarval activities Qi et al., 2008
Stigmast-22-en-3-one E. acoroides antifeedant, Qi et al., 2008
Stigmasta-5,22‑dien-3-O-b-D- E. acoroides antibacterial, antilarval activities Qi et al., 2008
glucopyranoside
Daucosterol E. acoroides antifeedant, Qi et al., 2008
Hexacosyl alcohol E. acoroides antibacterial, and antilarval activities Qi et al., 2008
p‑hydroxy-benzaldehyde E. acoroides antifeedant, Qi et al., 2008
Cholest-5-en3ß ol T. hemprichii Nichols and Johns, 1985
24-Methylcholesta-5,22E‑dien-3 ß ol T. hemprichii Nichols and Johns, 1985
24-Methylcholest-5-en-3ß ol T. hemprichii Nichols and Johns, 1985
24-Ethylcholesta-5,22E dien‑3 ß ol T. hemprichii Nichols and Johns, 1985
24-Ethylcholest-5-en-3 ß ol T. hemprichii Nichols and Johns, 1985
15,16-Epoxy‑ent-labda-8(17),13(16),14- R. maritima. Della Greca et al., 2000
trien-19-Ol
Methyl 15,16-epoxy‑ent-labda-8(17),13 R. maritima. Della Greca et al., 2000
(16),14-trien-19-oate
15,16-Epoxy‑ent-labda-8(17),13(16),14- R. maritima. Della Greca et al., 2000
trien-19-Al
15,16-Epoxy‑ent-labda-8(17),13(16),14- R. maritima. Della Greca et al., 2000
trien-19-ol acetate
Methyl-15,16-epoxy-12-oxo‑ent-labda-8 R. maritima. Della Greca et al., 2000
(17),13(16),14-trien-19-oate
ent-Labd-8(17),13E‑dien-15-ol R. maritima. Della Greca et al., 2000
13-Oxo-14,15-bis-nor‑ent-labd-8(17)- R. maritima. Della Greca et al., 2000
ene
D-galactan R. maritima. Aquino et al., 2005
Quercetin 3-O-b-D-galactopyranoside R. cirrhosa Enerstvedt et al., 2018
Quercetin 3-O-b-D-glucopyranoside R. cirrhosa
Quercetin3-O-b-d-(600 -O-malonyl) R. cirrhosa
galactopyranoside
Isorhamnetin 3-O-b-D-galactopyranoside R. cirrhosa
Isorhamnetin 3-O-b-D-glucopyranoside R. cirrhosa
Isorhamnetin3-O-b-D-(600 -O-malonyl) R. cirrhosa
galactopyranoside
60 ’’-acetyl syphonoside H. stipulacea Carbone et al., 2008
3,4,5-trihydroxybenzoic acid T. hemprichii Qi et al., 2012
5‑hydroxy-30 ,40 ,7-trimethoxyflavone T. hemprichii Qi et al., 2012
(E)
3-(4-methoxyphenyl)
2-propenoic T. hemprichii Qi et al., 2012
acid
40 ‑hydroxy-30 ,5,7-trimethoxyflavone T. hemprichii Qi et al., 2012
(S)‑methoxy-(30 ,50 -dimethoxy-40 - T. hemprichii Qi et al., 2012
hydroxyphenyl) ethanediol
Deoxycymodienol C. nodosa Antibacterial activity against methicillin- Kontiza et al., 2008
resistant (MRSA) strains
Isocymodiene C. nodosa Antibacterial activity against methicillin- Kontiza et al., 2008
resistant (MRSA) strains
Nodosol C. nodosa Antibacterial activity against methicillin- Kontiza et al., 2008
resistant (MRSA) strains
(1S*,2S*,3S*,7R*,8S*,9R*,11R*,12S*, C. nodosa Antibacterial activity against methicillin- Kontiza et al., 2008
14R*)
7‑bromo-tetradecahydro- resistant (MRSA) strains
12‑hydroxy-1-isopropyl-8,12-
dimethyl-4-methylenephenanthren-
9,14-yl diacetate

222
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al.
Table 3
Pharmacological activities of seagrass extracts.
Pharmacological activity Type of Extract
Antimicrobial Aqueous methanol extract
Ethanol extract
Hexane, chloroform,
methanol
Hexane, chloroform,
methanol
Chloroform, ethyl acetate,
ethanol and hexane
Ethyl acetate and hydro-
philic (water-soluble)
Petroleum ether, chloro-
form, ethyl acetate, ace-
tone, methanol water
Methanol extract
Petroleum ether, Chloroform S. Isoetifolium, H. ovalis, H.
fraction, butanol fraction
Methanol, hexane extract E. acoroides
Methanol/toluene (3:1)
extract
Ethanol, methanol extract
Methanol, ethanol, butanol,
water, acetone extracts
Ethanol, methanol, acetone,
dichloroethane
Diethyl ether, acetone,
methanol extracts
Ethyl acetate: methanol
(1:1) extract
Species
E. acoroides, T. hemprichii, H.
pinifolia, S. isoetifolium, C.
serrulata, C. rotundata
C. nodosa, R. cirrhosa
H. stipulaceae, H. pinifolia, C.
serrulata
H. stipulaceae, H. pinifolia, C.
serrulata
H. ovalis, C. serrulata, H.
pinifolia
E. acorioides, H. minor
C. serrulata, H. ovalis, Z.
capensis
S. isoetifolium
pinifolia
C. nodosa, P. oceanica, Z. noltii In-vitro
H. pinifolia, C. serrulata
C. rotundata
C. serrulata, S. isoetifolium
H. ovalis, H. pinifolia
T. testudinum, H. wrightii, S.
filiforme, H. decipiens, R.
maritima
223
South African Journal of Botany 137 (2021) 216
227
in vivo/ in vitro Model Reference
In-vitro S. aureus, Vibrio cholerae, Shi- Ragupathi Raja Kannan et al.,
gella dysentriae, Shigella 2012
bodii, S. paratyphi, P. aeru-
ginosa, K. pneumoniae
In-vitro Bacillus subtilis, Staphylococ- Abd El-Hady et al., 2007
cus aureus, E. coli, Pseudo-
monas eruginosa,
Aspergillus flavus, A. fumi-
gatus, and Alternaria
alternatium
In-vitro S. aureus, V. cholerae, Shigella Ragupathi Raja Kannan et al.,
dysentriae, Shigella bodii, S. 2010
paratyphi, P. aeruginosa, K.
pneumoniae
In-vitro Phytopathogens
Macro- Arumugam et al., 2010
phomina phaseolina, Colli-
totrichum capsici, Fusarium
sp., Aspergillus flavus, Pseu-
domonas aeruginosa
In-vitro E. coli, Enterococcus faecalis, Sangheetha & Asokan (2015)
Corynebacterium, Bacillus
subtilis Pseudomonas aeru-
ginosa, Klebsiella pneumo-
nia, Methicillin Sensitive
Staphylococcus aureus,
Methicillin Sensitive
Staphylococcus saphrophy-
ticus, Methicillin Sensitive
Staphylococcus epidermidis
In-vitro Lindra thalassiae, Dendry- Puglisi et al., 2007
phiella salina, Halophy-
tophthora spinosa,
Schizochytrium aggrega-
tum, Pseudoaltermonas
bacteriolytica
In-vitro Staphylococcus aureus, Bacil- Sreenathkumar et al., 2008
lus cereus, B. subtilis, E. coli,
Salmonella paratyphi, S.
Typhimurium, Micrococcus
luteus,
In-vitro Bacillus cereus, B. megate- Ravi Kumar et al., 2009
rium, Proteus vulgaris,
Streptococcus lactis, Staph-
ylococcus aureus, Aspergil-
lus niger, M. anisopliae
In-vitro Against 15 fouling bacteria Prabhadevi et al., 1997
In-vitro Bacillus subtilis, Staphylococ- Alam et al., 1994
cus aureus, Pseudomonas
aeruginosa, Klebsiella
pneumonia
E. coli, Bacillus subtilis, Can- Ballesteros et al., 1992
dida albicans, Aspergillus
niger
In-vitro Pseudomonas sp., Bacillus sp., Prabhakaran et al., 2012
Flavobacterium sp., Cyto-
phaga sp.
In-vitro V. cholerae, B. cereus, P. vul- Elizabeth Mani et al., 2012
garis, E. coli, S. dysenteriae,
S. paratyphi, Shigella sp., S.
mutants, S. aureus, P.
fluorescens
In-vitro Biofilm forming bacteria Mayavu et al., 2009
In-vitro Acenetobacter sp., S. typhii, Umamaheshwari et al., 2009
Micrococcus sp., Shigella
sonii, V. cholerae, Staphylo-
cocccus sp., Proteus vulga-
ris, P. mirabilis, P.
aeruginosa, Salmonella
paratyphi-B
In-vitro Fouling strains - Dendry- Ross et al., 2008
phiella salina, Lindra tha-
lassiae, Fusarium spp.
(continued)
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al.
Table 3 (Continued)
Pharmacological activity
Larvicidal
Cerium Binding Activity
Anti-inflammatory
Antiviral
Antimitotic
Cytotoxic
Antioxidant
Antidiabetic, antioxidant
and vasoprotective
Haemolytic activity
Type of Extract
Lipid and water soluble phe- H. ovalis, H. beccari, H.
nolic extracts
Ethanol extract and fractions C. serrulata
Ethanol extract and fractions C. serrulata
Methanol extract and
fractions
90% Aqueous methanol
Methanol extract H. ovalis
Petroleum ether, Chloroform S. Isoetifolium, H. ovalis
fraction
Aqueous ethanol S. isoetifolium, C. serrulata, H.
Pectin Z. marina, Phyllospadix
Methanol extract, EtOAC
fraction, Hexane fraction,
DCM fraction, Sub fraction
H5
Methanol extract
Isolated compounds
Methanol/toluene (3:1)
extract
70% Aqueous ethanol
70% Aqueous ethanol
Methanol/toluene (3:1)
extract
Methanol/toluene (3:1)
extract
Aqueous methanol extract
Pectin
Pectin
Pectin
Ethanol extract
Aqueous methanol
Methanol extract
Ethanol extract
Methanol, ethyl acetate, n-
hexane
Methanol extract
Ethyl acetate fraction
Aqueous methanol extract
Species
pinifolia
T. testudinum
C. rotundata
beccari
iwatensis
Z. japonica
H. ovalis
Thalassia testudinum
C. nodosa, P. oceanica, Z. noltii In-vivo
T. hemprichii
T. hemprichii, H. pinifolia
C. nodosa, P. oceanica, Z. noltii in-vivo
C. nodosa, P. oceanica, Z. noltii In-vivo
E. acoroides, T. hemprichii, H. In-vivo
pinifolia, S. isoetifolium, C.
serrulata, C. rotundata
Z. marina
Z. marina
Z. marina
E. acoroides
E.s acoroides, T. hemprichii,H.
pinifolia, Syringodium
isoetifolium
Halodule pinifolia, Halophila
ovalis, S. isoetifolium, T.
hemprichii, C. serrulata
E. acoroides, H. ovalis, H.
ovate, H. stipulacea, T. hem-
prichii, S. isoetifolium, C.
serrulata, H. pinifolia
T. hemprichii, C. rotundata, E.
acoroides, S. isoetifolium
H. ovalis
P. oceanica
E. acoroides, T. hemprichii, H.
pinifolia, S. isoetifolium, C.
serrulata, C. rotundata
224
South African Journal of Botany 137 (2021) 216
227
in vivo/ in vitro Model
In-vitro Against 9 human pathogens
In-vitro Fish pathogens
Bacillus
subtilis, Aeromonas hydro-
phila, Vibrio parahaemoly-
ticus, Serratia sp., V.
harveyii
In-vitro Poultry pathogens - Klebsi-
ella, E. coli, Staphylococcus
sp., Salmonella sp.
In-vitro Schizochytrium aggregatum
In-vitro 5 fouling strains - Bacillus
cereus, Bacillus circulans,
Bacillus pumilus, Pseudo-
monas vesicularis, Pseudo-
monas putida
In-vitro Human pathogens
In-vitro Brine shrimp bioassay
In-vitro Aedes aegypti larvae
In-vitro Can be used as removal of
radioisotopes from the
human body
In-vitro J774A.1 cells
In-vitro In-vitro
In-vivo HIV integrase inhibition test
herpes simplex virus, type I
(HSV) cultured in kidney
cells of monkey (CV-l),
vesicular stomatitis virus
(VSV) in kidney cells of
hamster (BAK)
In-vitro & In-vivo Encephalomyocarditis virus
(EMCV)
In-vitro Semliki forest virus (SFV) on
Vero cells
leukaemic cells of mice
(li210)
kidney cells of monkey (CV-
1)
Brine shrimp bioassay
In-vivo Lipid peroxidation
In-vitro Ferric reducing antioxidant
power
In-vivo Lipid peroxidation
In-vitro TAA, DPPH, FRAP assay
In-vitro TLC antioxidant assay
In-vitro Radical scavenging activity
In-vitro TAA, DPPH, FRAP assay
In-vitro DPPH radical assay
In-vitro Radical scavenging assay
In-vivo Glucose tolerance test, Liver
and kidney function tests,
In-vivo antioxidant
In-vivo Human erythrocyte
Reference
Balasubramaniam et al.,
2000
Ravikumar et al., 2011
Ravikumar et al., 2011
Jensen et al., 1998
Bhosale et al., 2002
Yuvaraj et al., 2012
Prabhadevi et al., 1998
Syed Ali et al., 2012
Khotimchenko et al., 2012
Hua et al., 2006
Yuvaraj et al., 2012
Rowley et al., 2002
Ballesteros et al., 1992
Premanathan et al., 1994
Premanathan et al., 1995
Ballesteros et al., 1992
Ballesteros et al., 1992
Ragupathi Raja Kannan et al.,
2012
Khasina et al., 2003
Kolenchenko et al., 2005
Sonina and. Khotimchenko,
2007
Ragupathi Raja Kannan et al.,
2010
Ragupathi Raja Kannan et al.,
2010
Athiperumalsamy et al.,
2010
Ragupathi Raja Kannan et al.,
2012
Santoso et al., 2012
Yuvaraj et al., 2012
Gokce and
Haznedaroglu, 2008
Ragupathi Raja Kannan et al.,
2012
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al. South African Journal of Botany 137 (2021) 216
227

Table 4
Seagrass species and their reported uses.

Plant species Family Uses References

Enhalus acoroides (L.f.) Royle
Enhalus acoroides (L.f.) Royle
Halophila ovalis (R.Br.) Hook.f.
Halophila gaudichaudii J. Kuo (syn. Halophila ovate
Gaud.)
Halophila stipulacea (Forssk.)Asch.
Syringodium isoetifolium (Asch.) Dandy
Thalassia hemprichii (Ehrenb. ex Solms) Asch.
Halodule pinifolia (Miki) Hartog
Halodule uninervis (Forssk.) Boiss.
Cymodocea serrulata (R.Br.) Asch. & Magnus
Cymodocea rotundata Asch. & Schweinf.
Hydrocharitaceae Rhizome and root juices consumed raw to treat Ramamurthy et al., 1991
seasickness. Rhizome (peeled of skin) con-
sumed fresh with cup of seawater to treat heart
conditions and low blood pressure. Rhizomes
consumed fresh to ease indigestion and hang-
over. Paste of fresh leaves used to treat many
kinds of skin diseases. Seeds, which taste like
almonds, are eaten by people or fed to goats
and sheep.
Hydrocharitaceae For the treatment of muscle aches, wounds, and Ratwanati et al., 2019
abdominal pain. Antidote against venomous
snake bites. The fruit can be used to alleviate
infectious diseases.
Hydrocharitaceae A handful of leaves is toasted with three drops of Ramamurthy et al., 1991; Kannan et al., 1999
sesame oil and consumed for three days to
treat iron deficiency. A leaf paste is mixed with
turmeric and applied to cure various skin ail-
ments, including burns and boils.
Hydrocharitaceae A handful of leaves is toasted with three drops of Newmaster et al., 2011
sesame oil and consumed for three days to
treat iron deficiency. A leaf paste is mixed with
turmeric and applied to cure various skin ail-
ments, including burns and boils.
Hydrocharitaceae A leaf paste is mixed with turmeric and applied to Newmaster et al., 2011
cure various skin ailments, including burns and
boils.
Potamogenetacea Fresh leaf juice consumed to ease acid reflux. Newmaster et al., 2011
Stems fed to goats.
Biomass used as green manure for garden.
Hydrocharitaceae Dried rhizome powder consumed to treat mental Newmaster et al., 2011
disorders. The same powder mixed with coco-
nut oil and applied on wounds. Biomass used as
green manure for gardens.
Potamogenetacea Stems are washed and used as goat feed. Newmaster et al., 2011
Potamogenetacea Stems are washed and used as goat feed. Newmaster et al., 2011
Potamogenetacea Branches and leaves used as feed for goats and Newmaster et al., 2011
pigs. Paste from leaves used to treat wounds.
Collected biomass used as green manure for
gardens.
Potamogenetacea Branches and leaves used as feed for goats and Newmaster et al., 2011
pigs. Paste from leaves used to treat wounds.
Collected biomass used as green manure for
gardens.

Cymodocea serrulata showed maximum activity in the methanol
extract of H. pinifolia (Kannan et al., 2010).
4.2. Antioxidant properties
Many natural antioxidants have been isolated from different
plants, such as oilseeds, a cereal crop, vegetables, spices and herbs
(Ramarathnam et al., 1995; Shon et al., 2003; Wettasinghe and Sha-
hidi, 1999). An immense number of marine flora and fauna have a
broad spectrum of interesting biological properties. Seagrasses are
rich in antioxidant compounds. Posidonia oceanica collected from the
western Mediterranean, showed increased GST (Glutathione Sulfo-
transferase) enzyme activity which is an essential indicator to antiox-
idant properties in plants (Rijstenbil et al., 1994; Hamountene et al.,
1995). In vivo antioxidant activity evaluation on male mice revealed
that the low etherified pectin isolated from Zostera marina normal-
izes the level of malonic dialdehyde, glutathione reductase and gluta-
thione peroxidase activity in the liver (Khasina et al., 2003). A
comparative study between the antioxidant activities of low-etheri-
fied pectin extracted from Z. marina and two antioxidative medicines
showed that zosterin and pectin manifested higher reducing activity
compared to other medicines (Kolenchenko et al., 2005). Pectin
extracted from Z. marina reduced the lead concentration in the liver,
decrease lipid peroxidation and normalize blood cholesterol levels in
rats (Sonina and Khotimchenko, 2007). Antidiabetic, antioxidant and
225
vasoprotective effects of P. oceanica extract (POE) were evaluated in
alloxan diabetic rats. The antidiabetic and vasoprotective effects of
POE was unrelated to its antioxidant properties (Gokce and Hazne-
daroglu, 2008). Thin layer chromatography analysis on methanolic
extracts of four seagrasses, namely E. acoroides, Thalassia hemprichii,
Halodule pinifolia and S. isoetifolium collected from Gulf of Mannar
followed by a 2, 2-diphenyl-1-picrylhydrazyl (DPPH) free radical test
on the isolated constituents to evaluate antioxidative properties
showed that antioxidant activities were comparatively stable in all
seagrasses investigated (Kannan et al., 2010b). Evaluation of antioxi-
dant activities of five seagrasses showed that the methanolic extracts
of Halophila ovalis had the highest antioxidant activity amongst the
species tested (Athiperumalsami et al., 2010).
4.3. Other medicinal properties
Seagrasses are an important natural biological source of diverse
metabolites with a huge array of bioactivities (Table 2 and 4). For
example, compounds with antifeedant and antilarval activities were
isolated and characterised from E. acoroides (Qi et al., 2008). Two dia-
rylheptanoids such as cymodienol, cymodiene (Kontiza et al., 2005)
and 3-keto steroids (Kontiza et al., 2006) were isolated from Cymodo-
cea nodosa and were reported as cytotoxic agents. The hexane sub-
fraction H5 of Zostera japonica has significant anti-inflammatory
activity.The major constituents were fatty acids (Hua et al., 2006).
D.H. Kim, M.F. Mahomoodally, N.B. Sadeer et al.
Thalassiolins A, B and C were isolated from Thalassia testudinum
(Rowley et al., 2002) as anti-HIV agents. Three compounds, namely
apigenin-7
0-b-D-glucoside, chrysoeriol and luteolin were isolated
from the ethyl acetate fraction of Z. marina. These compounds dis-
played antioxidative and anti-photoaging properties (Kim et al.,
2004). Free L-chiro-inositol (LCI), a very rare natural occurring cycli-
tol, was isolated from the flotsam of Syringodium filiforme
(Nuissier et al., 2008). Detritus of seagrasses Zostera noltii and Z.
marina was screened as a new source of zosteric acid (Achamlale
et al., 2009a) and rosmarinic acid (Achamlale et al., 2009b). Syphono-
sid was isolated from Halophila stipulacea (Carbone et al., 2008).
Chicoric acid was reported from leaves of Posidonia oceanica (Hazne-
daroglu and Zeybek, 2007) and S. filiforme (Nuissier et al., 2010).
From the foregoing account on biochemical and biomedical prop-
erties and phenolic acid content of seagrasses, it is evident that there
are gaps in information on these aspects of seagrasses entirely or at
least for many of the species. Hence, the present study was carried
out to highlights the areas where more research is needed.
5. Conclusion and future perspectives
This review showed that seagrasses produce many metabolites
with promising nutritional and bioactive potential such as antioxi-
dant, antimicrobial and anticancer. For example, Thalassiolins A, and
B were isolated from the Caribbean seagrass Thalassia testudinum and
Thalassia hemprichii exhibited potent cytotoxic and anti-HIV activi-
ties. Metabolites derived from seagrasses may provide useful leads
for future drug development. However, due to a scarcity of knowl-
edge on seagrasses, further consolidated investigation is necessary to
confirm these empirical findings. Furthermore, after conducting a
considerable literature search, a lack of enzymatic and in vivo studies
related to chronic diseases, namely diabetes, inflammatory diseases,
skin-related ailments was noted. Thus, for a better understanding of
the medicinal properties of seagrasses, it is of paramount importance
to conduct further in-depth investigations. After preliminary screen-
ing to identify promising natural products derived from seagrasses,
the compounds should be evaluated for safety and efficacy in animal
models and their mechanism and site of action determined before
testing these compounds in clinical trials and drug development.
Efforts should also focus on isolating minor metabolites of significant
pharmacological values using advanced technologies, namely NMR
micro cryogenic and capillary flow-probes and conducting biological
assays in smaller volume well plates and improving extraction meth-
ods.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
Acknowledgment
This paper was supported by Konkuk University in 2017.
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