Aquatic Microalgae As Potential Sources

Philippine Journal of Science
139 (1): 5-16, June 2010

ISSN 0031 - 7683
Aquatic Microalgae As Potential Sources

Of Uv-Screening Compounds
Maribel L. Dionisio-Sese
Institute of Biological Sciences, College of Arts and Sciences,
University of the Philippines Los Baos, College, Laguna 4031, Philippines
Microalgae are a polyphyletic and biochemically diverse assemblage of chlorophyll
a-containing microorganisms capable of oxygenic photosynthesis that are predominantly
found in aquatic environments with observed high levels of ultraviolet (UV) radiation.
Certain microalgae produce organic metabolites, such as sporopollenin, scytonemin and
mycosporine-like amino acids, to protect themselves from UV radiation while allowing visible
radiation involved in photosynthesis to pass through. Sporopollenin, an acetolysis-resistant
inert biopolymer usually observed in plant pollens and spores, was detected in the cell wall of
some UV-tolerant chlorophytes. Scytonemin, a yellow-brown lipid-soluble dimeric pigment,
was found in the extracellular polysaccharide sheath of some cyanobacteria. Mycosporinelike amino acids, which belong to a family of water-soluble compounds, were reported in
several free-living cyanobacteria, chlorophytes, haptophytes, diatoms, and dinoflagellates,
as well as in several marine invertebrate-microalgal symbiotic associations. Their capacity
to intercept UV radiation and dissipate its energy as heat without the formation of radical
intermediates makes these microalgal compounds potential sources of protection from UV
and photo-oxidative stress.
Key Words: microalgae, mycosporine-like amino acids, scytonemin, sporopollenin, UV-absorbing/

screening compounds, UV photoprotection
INTRODUCTION
Microalgae refers to a polyphyletic and biochemically
diverse assemblage of chlorophyll-a containing
microorganisms whose most distinct common
physiological attribute is their capability to undertake
oxygenic photosynthesis. They also share the characteristic
ability to thrive in aquatic environments wherein they are
regarded as the most important biomass producers on a
global scale.
High levels of visible and UV radiation in the tropical
region, particularly in the low-latitude tropics, occur due to
the shorter light path for sunlight in the region, the thinness
of the earths protective ozone layer near the equator, and
*Corresponding author: mldsese@yahoo.com
the high UV transparency of clear tropical ocean water. In

high latitudes, enhanced UV radiation due to stratospheric
ozone depletion is also a major stress factor for many
organisms. Ozone depletion has been observed in both
the Antarctic and Arctic regions, where ozone levels have
been reported to decline by as much as 50% during late
winter and early spring in the polar vortex (Smith et al.
1992; Von der Gathen et al. 1995).
UV radiation is customarily divided into three spectral
regions: UV-C (200-280 nm), UV-B (280-320 nm), and
UV-A (320-400 nm). UV-C, the most damaging spectral
region, is eliminated from the earths atmosphere through
absorption by ozone and other atmospheric gases. The
decreasing ozone concentration in the stratosphere, due
to the increasing concentrations of chlorofluocarbons,
5

Vol. 139 No. 1, June 2010
nitrogen dioxide and other pollutants in the atmosphere,

has led to increased penetration of UV-B, but not of UVA, on the surface of the earth (Kerr and McElroy 1993;
Madronich et al. 1998). Thus UV-B, which is deemed
deleterious, can penetrate biologically significant depths
(up to several meters) into the water column (Karentz
and Lutze 1990; Kuwahara et al. 2000), thereby affecting
aquatic ecosystems.
The detrimental effects of UV-B on primary producers are
varied, mediated primarily by damaging molecular targets
such as nucleic acids, proteins and pigments, and indirectly,
by producing reactive oxygen species. These can lead to
the inhibition of cell division and growth, affecting various
physiological and biochemical processes, such as motility,
orientation in motile organisms and photosynthesis, which
can consequently alter community structure and function.
UV-A can have a net damaging influence on photosynthesis
(Cullen et al. 1992). However, it is generally considered that
a more positive effect is exerted on physiology by inducing
photoreactivation or repair activities.
Sinha et al. (1998) identified the following four adaptation
strategies through which some organisms attempt to cope
with UV radiation:
(1) Availability of a number of DNA repair mechanisms,
such as photoreactivation and light-independent
nucleotide excision repair of DNA, and UV-A/blue
light-guided repair of the photosynthetic apparatus;
(2) Production of enzyme systems (e.g., superoxide
dismutase which reacts with and neutralizes the
highly toxic reactive oxygen species produced
by UV radiation) and induced formation of
quenching agents (e.g., carotenoids, especially
those in the xanthophyll cycle, which, in addition
to removing reactive oxygen species, can dissipate
excess excitation energy as heat or deactivate
excited chlorophyll, reducing photoinhibition of
photosynthesis);
(3) Behavioral modification to avoid exposure by
migrating to habitats with reduced UV exposure; and
(4) Production of UV-absorbing substances that screen
their cells from harmful UV radiation.
In terrestrial environments, where higher plants are the
foremost primary producers, several studies have shown
that harmful UV radiation in higher plants is absorbed by
epidermally located phenylpropanoids, mainly flavonoid
derivatives (Kootstra 1994). In aquatic environments,
where microalgae figure prominently, the presence and
role of UV-absorbing compounds like sporopollenin,
scytonemin, and mycosporine-like amino acids (MAAs)
have been established.
6
Dionisio-Sese ML: UV-Screemomg Compounds in Microalgae
Sporopollenin
Sporopollenin is an acetolysis-resistant inert biopolymer
possessing a complicated structure with aliphatic (mainly
isoprenoid) and aromatic components variably present.
It is found in the cell wall of some algae (Atkinson et
al. 1972; Pickett-Heaps and Staehelin 1975) and in plant
pollens and spores (Shaw and Yeadon 1964; Osthoff
and Wiermann 1987; Guilford et al. 1988). Xiong et al.
(1997) observed that UV-B tolerant chlorophyte species
of Characium terrestre, Coelastrum microporum, Enallax
coelastroides, Scenedesmus sp., Scotiella chlorelloidea,
Scotiellopsis rubescens, and Spongiochloris spongiosa
contain large amounts of sporopollenin. The biopolymer
was reported to be present also in Dunaliella salina
zygotes (Komaristaya and Gorbulin 2006).
Rozema et al. (2001) has demonstrated the UV-B
absorbing property of extracts containing sporopollenin.
The UV-B screening role of sporopollenin is evidenced
by the high UV-B optical density of the sporopollenins
isolated from UV-B tolerant chlorophytes and its
increase upon UV-B exposure. In Xiong et al. (1997),
the sporopollenin content (mean SE) increased from
0.730.12% to 0.850.14% (w/w) in Scenedesmus sp. and
from 0.58 0.10% to 0.730.09 % in Enallax coelastroides
upon UV-B exposure. Aside from its UV-B screening
role, the presence of sporopollenin in the cell wall of
Pediastrum duplex has been associated with the resistance
of this chlorophyte to microbial decomposition in natural
waters (Gunnison and Alexander 1975). The presence of
sporopollenin in the cell wall of Chlorella fusca also has
been associated to its resistance to extreme extraction
procedure (Atkinson et al. 1972).
Since sporopollenin is a non-toxic, safe natural material
that has the ability to absorb UV-B radiation and to bind
heavy metal (Arslan et al. 2004), its use in the cosmetic
treatment of skin against age-related/sun-related wrinkle
formation and as a chelating agent of ion-exchangers
in wastewater purification has been patented in the
United States in 2007 (http://www.patentgenius.com/
patent/7182965.html).
Scytonemin
Scytonemin is a yellow-brown lipid-soluble dimeric
pigment with a molecular mass of 544 kDa and a
structure based on indolic and phenolic subunits (Proteau
et al. 1993). It has an in vivo absorption maximum of
370 nm (in vitro at 386 nm) but purified scytonemin also
significantly absorbs at 252, 278, and 300 nm (Proteau
et al. 1993). Scytonemin can be readily extracted with
non-polar solvents such as acetone or ethyl acetate
and authenticated through spectrophotometry or
chromatography.

Vol. 139 No. 1, June 2010
Microalgal scytonemin appears restricted to cyanobacteria,

specifically in the extracellular polysaccharide sheath
of Chlorogloeopsis sp., Calothrix sp., Scytonema sp.,
Rivularia sp., and Nostoc commune (Sinha et al. 1998). It
is reportedly the most important UV-absorbing compound
in Lyngbya cf. aestuarii, where its area content seems to
follow the seasonal fluctuation of solar intensity (Karsten et
al. 1998). It was found to be the common pigment identified
in the extracellular sheath of more than 30 species of
cyanobacteria cultures and natural populations exposed to
intense solar radiation (Garcia-Pichel and Castenholz 1991).
In Chlorogloeopsis sp., UV-A exposure induced
photoinhibition and growth delay until substantial
concentrations of scytonemin have been synthesized
and deposited in the extracellular envelopes (GarciaPichel et al. 1992). In Calothrix sp., a correlation
between scytonemin content and photosynthetic
resistance to UV exposure has been demonstrated
(Brenowitz and Castenholz 1997; Dillon and Castenholz
2003). Garcia and Pichel (1991) strongly suggested
that scytonemin production constitutes an adaptive
strategy of photoprotection against UV irradiation.
There are several studies indicating that the incident
UV-A radiation entering the cells may be reduced
by around 90% due to the presence of scytonemin in
the cyanobacterial sheaths (Garcia and Pichel 1991;
Garcia-Pichel et al. 1992; Brenowitz and Castenholz
1997). Further, Dillon and Castenholz (2002) have
demonstrated that scytonemin protects cyanobacteria
against UV-C radiation, even suggesting that this
pigment may have evolved during the Precambrian era
and made possible the colonization of exposed shallow
waters and terrestrial habitats by cyanobacteria or their
oxygenic ancestors.
Scytonemin is suspected to be synthesized from metabolites
of aromatic amino acid biosynthesis. Molecular genetics
and genomic analysis indicate that tryptophan and
tyrosine are the likely biosynthetic precursors of
scytonemin (Soule et al. 2007). Aside from the induction
of scytonemin synthesis by high photon fluence rate and
UV-A (Garcia-Pichel and Castenholz 1991), Dillon et al.
(2002) reported that, in Chroococcidiopsis sp., an increase
in both temperature and oxidative stress in combination
with UV-A, gave a synergistic effect on high production
of scytonemin. Periodic desiccation also enhanced the
synthesis of scytonemin in Nostoc punctiforme (Fleming
and Castenholz 2007). The high stability of scytonemin
against stressors such as temperature and UV-A radiation
and its capability to undertake UV screening even after
prolonged physiological inactivity such as desiccation
make it a strong candidate for use as a natural UVscreening compound for humans (Sinha and Hder 2008).
Aside from its UV-absorbing property, scytonemin was

found to have anti-inflammatory and anti-proliferative
capabilities through its inhibition of human polo-like
kinase (Stevenson et al. 2002a, 2002b). In 2002, the
United States has patented methods of using scytonemin
in treating disorders associated with cell cycle progression,
cell proliferation, kinase activity, tissue hyperplasia or
angiogenesis, as in cancer or inflammatory diseases
(http://www.patentgenius.com/patent/6495586.html).
Mycosporine-Like Amino Acids
Mycosporine-like amino acids or MAAs are watersoluble substances with 310-360 nm absorption maxima
and average molecular weight of 300 kDa (Nakamura
et al. 1982). They are characterized by a cyclohexenone
or cyclohexenimine chromophore conjugated with the
nitrogen substituent of an amino acid or its imino alcohol.
The aromatic chromophore mycosporine was first isolated
from fungi, hence the name. Its production in fungi
coincides with UV-stimulated reproduction stage (Leach
1965). Unlike sporopollenin and scytonemin, which
are found in the cell wall or extracellular sheath of the
microalgae, MAAs in microalgae are mostly intracellular.
Table 1 lists the MAAs found in various organisms to date
(Sinha and Hder 2008). The variations in the absorption
spectra of the MAAs are due to the different attached side
groups and nitrogen substituents.
Table 1. Some identified mycosporine-like amino acids and their
corresponding wavelengths of maximum absorption.
Mycosporine-like amino acids
Absorption maximum
(max)
Asterina
330 nm
Euhalothece
362 nm
Mycosporine-glutamic acid:glycine
330 nm
Mycosporine-glycine
310 nm
Mycosporine-2-glycine
334 nm
Mycosporine-glycine:valine
335 nm
Mycosporine-methylamine-serine
327 nm
Mycosporine-methylamine-threonine
327 nm
Mycosporine-taurine
309 nm
Palythene
360 nm
Palythenic acid
337 nm
Palythine
320 nm
Palythine-serine
320 nm
Palythine-serine-sulfate
320 nm
Palythinol
332 nm
Porphyra
334 nm
Shinorine
334 nm
Usujirene
357 nm

Vol. 139 No. 1, June 2010
Table 2. Species of cyanobacteria and their mycosporine-like amino acids.

Species
Identified MAAs
Reference
Anabaena doliolum
mycosporineglycine, porphyra, shinorine
Singh et al. (2008b)
Anabaena sp.
shinorine
Sinha et al. (1999)
Anabaena variabilis PCC 7937
shinorine
Singh et al. (2008a)
Chlorogloeopsis PCC 6912
mycosporine-glycine, shinorine
Portwich and Garcia-Pichel (1999)
Euhalothece sp.
euhalothece, mycosporine-2-glycine
Kedar et al. (2002), Volkmann et al. (2006)
Gloeocapsa sp.
mycosporine-glycine, asterina, palythinol, shinorine
Sommaruga and Garcia-Pichel (1999)
Microcoleus chthonoplastes
shinorine
Karsten (2002)
Microcystis aeruginosa
porphyra, shinorine
Liu et al. (2004)
Nodularia baltica
porphyra, shinorine
Sinha et al. (2003b)
Nodularia harveyana
porphyra, shinorine
Nodularia spumigena
porphyra, shinorine
Nostoc commune
shinorine
Sinha et al. (2001, 2003a)
Scytonema sp.
shinorine
Sinha et al. (2001)
Synechocystis sp. PCC 6803
mycosporine-taurine, usujirene-like
Zhang et al. (2007)
MAAs are usually extracted using methanol and separated

through high-performance liquid chromatography. They can
be identified and quantified based on their absorption peaks
and retention times and co-chromatography with available
standards for validation (Nakamura et al. 1982). In the
absence of well-defined or commercially available standards,
electrospray ionization mass spectrometry coupled with
liquid chromatography also has been used to examine MAA
diversity and analyze MAAs (Whitehead and Hedges 2002).
A survey of several strains of cyanobacteria by GarciaPichel and Castenholz (1993) established the presence of
one or more water-soluble UV-absorbing MAAs in these
microorganisms. Although the specific MAA contents
varied among strains, the quantities of MAAs found were
invariably higher when cultures were grown with UV
radiation than when UV was absent. Since MAAs are
photoinducible, their presence may also explain the ability
of some cyanobacteria to develop and maintain surface
blooms even in the presence of high solar irradiance,
including UV radiation (Liu et al. 2004). Table 2 lists
the MAAs identified in several species of cyanobacteria.
The presence of several MAAs in a species also was
observed in different chlorophytes, haptophytes, diatoms,
and dinoflagellates (Table 3) grown in cultures or collected
in a broad variety of aquatic habitats, from the tropics to
the polar regions. Although the specific MAA contents
varied among species, they also were found to be generally
more when the microalgal species were grown with UV
radiation than when without it. Thus, high contents of
8
MAA in the cells correlate with increased resistance to

UV radiation. Individually, MAAs have wide absorption
bands at about 40 nm (Sinha et al. 1998). Hence, the
cumulative effect of having several MAAs with different
absorption maxima greatly benefits an organism since
the UV-filtering capacity is further broadened, increasing
protection across a wider range of wavelengths. On the
other hand, the varying capacities of different microalgal
species to produce UV-screening compounds could
potentially result in changing species dominance in
the community. Species producing high concentrations
of UV-screening compounds may form dense surface
blooms (Liu et al. 2004) thereby affecting the structure
and dynamics of aquatic ecosystems and trophic flow.
MAAs also have been reported in several microalgaeinvertebrate (sea anemone, coral, ascidian) symbiotic
associations such as those in Table 4. Such photoautotrophic
symbioses allow for the beneficial exchange of nutrients
between the microalgae and their animal hosts.
In a more extensive survey of MAAs in 54 species of
symbiotic cnidarians that included hydrozoan corals,
anemones, gorgonians and scleractinian corals, Banaszak
et al. (2006) found that the symbiont fractions, under
natural conditions, synthesized one to four MAAs. They
further suggested that Symbiodinium sp. is restricted to
producing five MAAs with a defined order of appearance:
first, mycosporine-glycine; followed by shinorine (in
one case mycosporine-2-glycine); followed by porphyra;
finally, by palythine.

Vol. 139 No. 1, June 2010
Table 3. Microalgal groups/species and their mycosporine-like amino acids.

Microalgal Group/Species
Identified MAAs
Reference
Chlorophytes
Ankistrodesmus spiralis
asterina, shinorine
Xiong et al. (1999)
Chlorella minutissima
asterina, shinorine
Xiong et al. (1999)
Chlorella sorokiniana
palythine, porphyra, shinorine
Xiong et al. (1999)
Dunaliella tertiolecta
mycosporine-glycine
Hannach and Sigleo (1998)
Enallax coelastroides
Xiong et al. (1999)
Pseudococcomyxa sp.
mycosporine-glycine, palythine
Xiong et al. (1999)
Pyramimonas parkeae
mycosporine-glycine
Scenedesmus sp.
asterina, palythine, porphyra, shinorine
Xiong et al. (1999)
Scotiella chlorelloidea
asterina, palythine, porphyra, shinorine
Xiong et al. (1999)
Isochrysis sp.
mycosporine-glycine
Pavlova gyrans
mycosporine-glycine, porphyra/shinorine
Chaetoceros sp.
porphyra, shinorine
Riegger and Robinson (1997)
Corethron criophilum
porphyra, shinorine
Helbling et al. (1996)
Cosinodiscus centralis
porphyra, shinorine
Thalassiosira tumida
porphyra, shinorine
Porosira glacialis
porphyra, shinorine
Porosira pseudodenticulata
mycosporine-glycine, porphyra, shinorine
Proboscia inermis
mycosporine-glycine
Stellarima microtrias
porphyra, shinorine
Thalassiosira antarctica
porphyra, shinorine
Thalassiosira tumida
porphyra, shinorine
Thalassiosira sp.
porphyra, shinorine
Helbling et al. (1996)
Thalassiosira weissflogii
mycosporine-glycine
Haptophytes
Diatoms
Dinoflagellates
Alexandrium catenella
Alexandrium excavatum
Alexandrium minutum
Alexandrium tamarense
Amphidinium carterae
Gymnodinium catenatum
Gymnodinium sanguineum
Gyrodinium dorsum
Heterocapsa sp.
Lingulodinium polyedra
Prorocentrum micans
Prorocentrum minimum
Scrippsiella sweeneyae
mycosporine-methylamine-serine, mycosporineglycine, palythene, palythenic acid, palythine,

palythinol, porphyra, shinorine, usujirene
palythene, porphyra, shinorine, usujirene
mycosporine-glycine, palythene, palythenic
acid, palythine, palythinol, porphyra, shinorine,
usujirene
asterina, mycosporine-glycine, palythene, palythenic acid, palythine, palythinol, porphyra,
shinorine, usujirene
mycosporine-glycine
Carreto et al. (2001)

mycosporine-glycine, porphyra, shinorine

mycosporine-glycine, palythine, paythene,
porphyra
Jeffrey et al. (1999)
mycosporine-2-glycine, shinorine, palythinol

mycosporine-glycine:valine, palythene, palythine, palythinol, porphyra
asterina, mycosporine-glycine, porphyra, shinorine
palythene, shinorine
mycosporine-glycine, palythene, palythine,
porphyra, shinorine
Montero and Lubin (2003)
Neale et al. (1998)

Klisch et al. (2001)
Vernet and Whitehead (1996)
Lesser (1996)
Sinha et al. (1999)
Taira et al. (2004)

Vol. 139 No. 1, June 2010
Table 4. Mycosporine-like amino acids in various microalga-host symbiotic associations.

Microalga-Host Association
Identified MAAs
Symbiodinium sp.-Cassiopea xamachana
mycosporine-glycine, mycosporineBanaszak and Trench (1995)

taurine, porphyra, shinorine
Symbiodinium sp.-Anthopleura elegantissima
mycosporine-glycine, mycosporineStochaj et al. (1994)

Symbiodinium sp.-Phyllodiscus semoni
mycosporine-glycine, mycosporineShick et al. (1991)

Symbiodinium sp.-Clavularia sp.
asterina, palythene, palythine
Symbiodinium sp.-Acropora microphthalma
mycosporine-glycine, mycosporineShick et al. (1995)

Prochloron sp.-Lissoclinum patella
mycosporine-glycine, palythine, shinorine Dionisio-Sese et al. (1997)
Among the microalga-host symbiotic associations, the

Symbiodinium-coral and Prochloron-ascidian symbioses
generate substantial interest. Coral reefs are highly
productive and support dense populations of marine
organisms. They have been receiving increased scientific
and media attention due to the increasing incidence of
bleaching on a global scale. Considerable attention is
drawn to the Prochloron-ascidian symbiosis because
of its unique ecological distribution and microsymbiont
biochemical characteristic. Ecologically, Prochloron is
exclusively associated with tropical didemnid ascidians.
Biochemically, it is a prokaryote although it lacks the
phycobiliproteins typical of the cyanobacteria. Instead,
in addition to chlorophyll a, it possesses chlorophyll b
and light harvesting a/b protein, which is characteristic of
eukaryotic chlorophytes (Lewin 1975, 1981). Prochloron
also shows carbonic anhydrase inhibition values similar
to unicellular cyanobacterium and chloroplasts of green
algae and higher plants (Dionisio-Sese et al. 1993).
Thus, it could be placed as a possible evolutionary link
between cyanobacteria and chlorophytes (Dionisio-Sese
1996). Small subunit-ribosomal RNA analysis, however,
seems to have established that prochlorophytes are more
related with cyanobacteria (Urbach et al. 1992) and that
their chlorophyll a/b-binding proteins are unrelated to
those from plastids of green algae and higher plants (La
Roche et al. 1996).
In the Prochloron-ascidian symbiosis, Dionisio-Sese et al.
(1997) correlated the predominance of MAAs in the upper
layer of the host tissue with its photoprotective role against
UV radiation. Ultraviolet radiation severely inhibited
Prochloron photosynthesis in isolation but not in vivo.
This protection was provided by the upper tunic covering
the ascidian colony, which contains MAAs (Dionisio-Sese
et al. 1997, 2001). Since the closely related mycosporines
in fungi are synthesized via the shikimic acid pathway
(Favre-Bonvin et al. 1987), which does not occur in
animals but only in bacteria, fungi, algae, and plants, it
10
Reference
Shick et al. (1991)
can be inferred that the MAAs may have been produced

by the symbiont and transferred to the host where it
underwent structural modification. Evidence exists that
the primary MAA mycosporine-glycine, synthesized from
the precursor 3-dehydroquinone via gadusols (Shick and
Dunlap 2002), can be transformed into other secondary
MAAs (Callone et al. 2006). On the other hand, alternative
origins of these compounds in animals, either dietary or
de novo synthesis, can also occur (Stochaj et al. 1994;
Carroll and Shick 1996; Newman et al. 2000). Thus,
MAAs appear capable of providing protection from UV
radiation not only to their producers but also to their
symbiotic partners via translocation or to their primary
and secondary consumers through accumulation via the
food chain. This seems supported by the observation that
MAAs are also highly resistant against abiotic stressors
such as temperature, various solvents, and pH (Grniger
and Hder 2000).
There are strong evidences that the primary biological
function of MAAs in various organisms is as UVsunscreens, protecting cells from UV-induced damage
by absorbing incoming UV radiation (Garcia-Pichel et
al. 1993; Dionisio-Sese et al. 1997; Neale et al. 1998;
Klisch et al. 2001). Since MAAs prevent three out of ten
photons from hitting cytoplasmic targets (Sinha and Hder
2002), inhibition of photosynthesis and growth, pigment
destruction, membrane disruption, enzyme inactivation,
DNA damage, and, finally, cell death are averted. It
has been established that MAAs offer protection from
UV and photooxidative stress by intercepting 310-360
nm UV wavelengths and dissipating the energy as heat
without producing reactive oxygen species (Shick 1993;
Conde et al. 2000). Their photoprotective role also is
supported by their photostability in both fresh and sea
water in the presence of photosensitizers (Whitehead and
Hedges 2005). Such ideal characteristics of MAAs as
photoprotectants have led to the development of synthetic
simple MAA analogues in Australia, some of which have

Vol. 139 No. 1, June 2010
been tested for use in human skin-care and cosmetic

products (Dunlap et al. 1998). In 2004, the United States
issued a patent for compounds having MAAs, in addition
to scytonemin and carotenoids, as active sunscreen agents
that can be used as solar protection for humans (http://
www.patentgenius.com/patent/6787147.html).
The intracellular location of MAAs suggests that they
perform other biological functions aside from serving
as UV-screens. Mycosporine-glycine has been reported
to have antioxidant activity against peroxyl-radical
initiated lipid peroxidation (Dunlap and Yamamoto
1995; Suh et al. 2003). Oren (1997) reported that MAAs
may also function as osmolytes or osmotic regulators
especially in cyanobacteria. In Chlorogloeopsis PCC
6912, mycosporine-glycine production could be induced
by osmotic stress (Portwich and Garcia-Pichel 1999).
Shinorine synthesis has been induced by salt stress in
Microcoleus chthonoplastes (Karsten 2002) and Anabaena
variabilis PCC 7937 (Singh et al. 2008a). These indicate
possible double functions of MAAs as organic osmolytes
and photoprotectants in these organisms.
Carotenoids, although non-UV absorbing pigments, can
be included as components of sunscreen products since
they also play major roles in photoprotection by promoting
heat dissipation of excess light energy and facilitating
removal of reactive oxygen species. The accumulation
of carotenoids under excessive light is typically observed
in surface populations of cyanobacteria, chlorophytes,
and dinoflagellates (Pearl et al. 1983; Ben-Amotz et al.
1989; Vernet et al. 1989). Photoprotection by carotenoids
is brought about specifically by the xanthophyll cycle
which promotes dissipation of excess excitation energy
as heat in the light-harvesting centers of photosynthesis.
Deactivation of excited chlorophyll is afforded by the
conversion of the xanthophyll pigments violaxanthin
to zeaxanthin in cyanobacteria and chlorophytes or of
diadinoxanthin to diatoxanthin in dinoflagellates. In
addition to carotenoids, Araoz and Hder (1997) suggested
phycobiliproteins to be effective screening agents in
cyanobacteria since they absorb in the UV-B range, plus
they form a peripheral layer around the central part of the
cell containing the DNA. They also showed increased
synthesis of phycobiliprotein pigments under mild UV-B
stress, simultaneous with its destruction.
CONCLUSION
Much information on UV-absorbing compounds
(sporopollenin, scytonemin, and MAAs) from microalgae
that predominantly thrive in tropical and polar waters have
been established over the past decades. These compounds
play significant roles in the growth and survival of

microalgae in natural habitats, particularly in coping with
UV radiation. Aside from their UV-screening capacity,
they also seem to perform other physiological functions in
the microalgae that produce them. The resistance of some
chlorophytes to extraction and microbial decomposition
in natural waters is being attributed to the presence of
sporopollenin in their cell walls. Scytonemin has been
found to have anti-inflammatory and anti-proliferative
capabilities. It is considered to have an important
ecophysiological role in the evolutionary history of
cyanobacteria.
MAAs have been reported to act as antioxidants and
as osmotic regulators. The photostable MAAs, with
broad absorption maxima between 310-360 nm, provide
photoprotection by effectively dissipating absorbed
radiation as heat without producing reactive oxygen
species, shielding the organism from UV-induced
damages. Since MAAs are found also in organisms
lacking the shikimate pathway, MAAs provide UVprotection not only to their producers but also to their
primary and secondary consumers through the food chain.
Thus, the presence of MAAs can enhance the primary
productivity of an aquatic ecosystem by facilitating the
growth and survival of the microalgae even under intense
UV exposure. The impact of this increased biomass
productivity would be reflected through all levels of the
intricate food web, resulting ultimately in increased food
production for humans.
As safe and natural biochemicals, microalgal UVscreening compounds clearly present potential
commercial uses that have yet to be fully explored,
particularly in the Philippines. The United States has
issued a patent on methods of using sporopollenin for
cosmetic purposes and as chelating agents for wastewater
treatment. The development of synthetic simple
MAA analogues for cosmetic products has long been
underway in Australia whereas sunscreens containing
photoprotective compounds like scytonemin and MAAs
are already available in the United States.
The multi-billion dollar global skin care and cosmetic
business might be already seriously looking at microalgal
UV-screening compounds as its wave of the future. The
Philippines multi-billion peso skin care and cosmetic
industry, including its alternative health care sector, can
substantially do well in tapping the commercial potentials
of microalgal UV-screening compounds this early. It does
not even need to follow the synthetic route continental
countries seem to be taking as the countrys archipelagic
nature and its vast territorial waters offer a strong strategic
advantage in the mass production of aquatic microalgae.
11

Vol. 139 No. 1, June 2010
Research investment can hasten the fine-tuning of microalgal

UV-screeners industrial applications. Continuing studies
of microalgae may reveal still unknown UV-screening
compounds or new microalgal species that produce more
of the presently known UV-screeners.
REFERENCES
ARAOZ R, HDER DP. 1997. Ultraviolet radiation induces
both degradation and synthesis of phycobilisomes in
Nostoc sp.: a spectroscopic and biochemical approach.
FEMS Microbiol Ecol 23:301-313.
ARSLAN M, TEMOCIN Z, YIGITOGLU M. 2004.
Removal of cadmium (II) from aqueous solutions using
sporopollenin. Fresenia Env Bull 13: 616-619.
ATKINSON AW, GUNNING BES, JOHN PCL. 1972.
Sporopollenin in the cell wall of Chlorella and other
algae: Ultrastructure, chemistry and incorporation of
14
C-acetate, studied in synchronous cultures. Planta
107:1-32.
BANASZAK AT, TRENCH RK. 1995. Effects of
ultraviolet (UV) radiation on marine microalgalinvertebrate symbioses. II. The synthesis of
mycosporine-like amino acids in response to exposure
to UV in Anthopleura elegantissima and Cassiopea
xamachana. J Exptl Mar Biol Ecol 194:233-250.
BANASZAK AT, SANTOS MGB, LAJEUNESSE TC,
LESSER MP. 2006. The distribution of mycosporinelike amino acids (MAAs) and the phylogenetic identity
of symbiotic dinoflagellates in cnidarian hosts from the
Mexican Caribbean. J Exptl Mar Biol Ecol 337:131-146.
BEN-AMOTZ A, SHAISH A, AVRON M. 1989. Mode
of action of the massively accumulated -carotene
of Dunaliella bardawil in protecting the alga against
damage by excessive radiation. Plant Physiol
91:1040-1043.
BRENOWITZ S, CASTENHOLZ RW. 1997. Longterm effects of UV and visible irradiance on natural
populations of a scytonemin-containing cyanobacterium
(Calothrix sp.). FEMS Microbial Ecol 24:343-352.
CALLONE AI, CARIGNAN M, MONTOYA NG,
CARRETO JI. 2006. Biotransformation of mycosporine
like amino acids (MAAs) in the toxic dinoflagellate
Alexandrium tamarense. J Photochem Photobiol B
Biol: 84:204-212.
CARRETO JI, CARIGNAN MO, DALEO G, DE
MARCO SG. 1990. Occurrence of mycosporine-like
amino acids in the red tide dinoflagellate Alexandrium
12
excavatum: UV-protective compounds? J Plankton Res

12:909-921.
CARRETO JI, CARIGNAN MO, MONTOYA NG. 2001.
Comparative studies on mycosporine-like amino
acids, paralytic shellfish toxins and pigment profiles
of the toxic dinoflagellates Alexandrium tamarense,
A. catenella and A.minutum. Mar Ecol Prog Ser 223:
49-60.
CARROLL AK, SHICK JM. 1996. Dietary accumulation
of UV-absorbing mycosporine-like amino acids
(MAAs) by the green sea urchin (Strongylocentrotus
droebachiensis). Mar Biol 124:561-569.
CONDE FR, CHURIO MS, PREVITALI CM. 2000.
The photoprotector mechanism of mycosporine-like
amino acids. Excited state properties and photostability
of porphyra-334 in aqueous solution. J Photochem
Photobiol B Biol 56:139-144.
CULLEN JJ, NEALE PJ, LESSER MP. 1992. Biological
weighting function for the inhibition of phytoplankton
photosynthesis by ultraviolet radiation. Science
258:646-650.
DILLON JG, CASTENHOLZ RW. 2002. Scytonemin, a
cyanobacterial sheath pigment, protects against UVC
radiation: implications for early photosynthetic life. J
Phycol 35:673-681.
DILLON JG, CASTENHOLZ RW. 2003. The synthesis
of the UV-screening pigment, scytonemin, and
photosynthetic performance in isolates from closely
related natural populations of cyanobacteria (Calothrix
sp.). Env Microbiol 5:484-491.
DILLON JG, TATSUMI CM, TANDINGAN PG,
CASTENHOLZ RW. 2002. Effect of environmental
factors on the synthesis of scytonemin, a UV-screening
pigment, in a cyanobacterium (Chroococcidiopsis sp.).
Arch Microbiol 177:322:331.
DIONISIO-SESE ML. 1996. Carbonic anhydrase: Its
physiological and evolutionary significance in the
marine symbiont Prochloron. Phil J Sci 125:241-254.
DIONISIO-SESE ML, ISHIKURA M, MARUYAMA
T, MIYACHI S. 1997. UV-absorbing substances in
the tunic of a colonial ascidian protect its symbiont,
Prochloron sp., from damage by UV-B radiation. Mar
Biol 128:455-461.
DIONISIO-SESE ML, MARUYAMA T, MIYACHI S.
2001. Photosynthesis of Prochloron as affected by
environmental factors. Mar Biotechnol 3:74-79.
DIONISIO-SESE ML, SHIMADA A, MARUYAMA T,
MIYACHI S. 1993. Carbonic anhydrase activity of

Vol. 139 No. 1, June 2010
Prochloron sp. isolated from an ascidian host. Arch

Microbiol 159:1-5.
DUNLAP WC, YAMAMOTO Y. 1995. Small-molecule
antioxidants in marine organisms: antioxidant activity
of mycosporine-glycine. Comp Biochem Physiol
112B:105-114.
DUNLAP WC, CHALKER BE, BANDARANAYAKE
WM, WU WON JJ. 1998. Natures sunscreen from
the Great Barrier Reef, Australia. Int J Cosmetic Sci
20: 41-51.
FAVRE-BONVIN J, BERNILLON J, SALIN N, ARPIN
N. 1987. Biosynthesis of mycosporines: mycosporine
glutaminol in Trichothecium roseum. Phytochem
26:2509-2514.
FLEMING ED, CASTENHOLZ RW. 2007. E f f e c t s
of periodic desiccation on the synthesis of the UVscreening compound, scytonemin, in cyanobacteria.
Environ Microbiol 9:1448-1455.
GARCIA-PICHEL F, CASTENHOLZ RW. 1991.
Characterization and biological implications of
scytonemin, a cyanobacterial sheath pigment. J Phycol
27:395-409.
GARCIA-PICHEL F, CASTENHOLZ RW. 1993. Occurrence
of UV-absorbing, mycosporine-like compounds among
cyanobacterial isolates and an estimate of their screening
capacity. Appl Env Microbiol 59:163-169.
GARCIA-PICHEL F, SHERRY ND, CASTENHOLZ RW.
1992. Evidence for an ultraviolet sunscreen role of
the extracellular pigment scytonemin in the terrestrial
cyanobacterium Chlorogloeopsis sp. Photochem
Photobiol 56:17-23.
GARCIA-PICHEL F, WINGARD CE, CASTENHOLZ
RW. 1993. Evidence regarding the UV sunscreen role of
a mycosporine-like compound in the cyanobacterium
Gloeocapsa sp. Appl Env Microbiol 59:170-176.
GRNIGER A, HDER DP. 2000. Stability of
mycosporine-like amino acids. Recent Res Devel
Photochem Photobiol 4:247-252.
GUILFORD WJ, SCHNEIDER DM, LABOVITZ J,
OPELLA SJ. 1988. High resolution solid state 13C
NMR spectroscopy of sporopollenins from different
plant taxa. Plant Physiol 86:134-136.
GUNNISON D, ALEXANDER M. 1975. Basis for the
resistance of several algae to microbial decomposition.
Appl Microbiol 29: 729-738.
HANNACH G, SIGLEO AC. 1998. Photoinduction of
UV-absorbing compounds in six species of marine
phytoplankton. Mar Ecol Prog Ser 174:207-222.
H E L B L I N G E W, C H A L K E R B E , D U N L A P
WC, OSMUND HH, VILLAFANE VE. 1996.
Photoacclimation of Antarctic diatoms to solar
ultraviolet radiation. J Exptl Mar Biol Ecol 204:85-101.
http://www.patentgenius.com/patent/6495586.html
referred on 14 February 2009.
referred on 01 March 2009.
referred on 21 February 2009.
JEFFREY SW, MAC TAVISH HS, DUNLAP WC,
VESK M, GROENEWOUD K. 1999. Occurrence of
UVA- and UVB-absorbing compounds in 152 species
(206 strains) of marine microalgae. Mar Ecol Prog
Ser 189-35-51.
KARENTZ D, LUTZE H. 1990. Evaluation of
biologically harmful ultraviolet radiation in Antarctica
with a biological dosimeter designed for aquatic
environments. Limnol Ocenogr 35:549-561.
KARSTEN U. 2002. Effects of salinity and ultraviolet
radiation on the concentration of mycosporinelike amino acids in various isolates of the benthic
cyanobacterium Microcoleus chthonoplastes. Phycol
Res 50:129-134.
KARSTEN U, MAIER J, GARCIA-PICHEL F.
1998. Seasonality in UV-absorbing compounds of
cyanobacterial mat communities from an intertidal
mangrove flat. Aquatic Microbial Ecol 16:37-44
KEDAR L, KASHMAN Y, OREN A. 2002. Mycosporine2-glycine is the major mycosporine-like amino acid in a
unicellular cyanobacterium (Euhalothece sp.) isolated
from a gypsum crust in a hypersaline saltern pond.
FEMS Microbiol Lett 208:233-237
KERR JB, MCELROY CT. 1993. Evidence for large
upward trends of ultraviolet-B radiation linked to ozone
depletion. Science 262:1032-1034.
KLISCH M, SINHA RP, RICHTER PR, HDER DP.
2001. Mycosporine-like amino acids (MAAs) protect
against UV-B induced damage in Gyrodinium dorsum
Kofoid. J Plant Physiol 158:1449-1454.
KOMARISTAYA VP, GORBULIN OS. 2006.
Sporopollenin in the composition of cell walls of
Dunalliela salina Teod. (Chlorophyta) zygotes. Int J
Algae 8:43-52.
KOOTSTRA A. 1994. Protection from UV-B induced
DNA damage by flavonoids. Plant Mol Biol 26:771774.
13

Vol. 139 No. 1, June 2010
KUWAHARA VS, TODA T, HAMASAKI K, KIKUCHI

T, TAGUCHI S. 2000. Variability in the relative
penetration of ultraviolet radiation to photosynthetically
active radiation in temperate coastal waters, Japan. J
Oceanogr 56:399-408.
LA ROCHE J, VAN DER STAAY GWM, PARTENSKY
F, DUCRET A, AEBERSOLD R, LI R, GOLDEN
SS, HILLER RG, WRENCH PM, LARKUM AWD,
GREEN BR. 1996. Independent evolution of the
chlorophyte and green light chlorophyll a/b lightharvesting proteins. Proc Natl Acad Sci 93:1524415248.
LEACH CM. 1965. Ultraviolet-absorbing substances
associated with light-induced sporulation in fungi. Can
J Bot 43:185-200.
LESSER MP. 1996. Acclimation of phytoplankton to
UV-B radiation: oxidative stress and photoinhibition
of photosynthesis are not prevented by UV-absorbing
compounds in the dinoflagellate Prorocentrum micans.
Mar Ecol Prog Ser 132:287-297.
LEWIN RA. 1975. A marine Synechocystis (Cyanophyta,
Chlorococcales) epizoic on ascidians. Phycologia
14:153-160.
LEWIN RA. 1981. The prochlorophytes. In: Starr MP,
Stolp H, Truper HG, Balows A, Schlegel HG, editors.
The Prokaryotes. Berlin: Springer Verlag. p. 257-266.
LIU A, HADER DP, SOMMARUGA R. 2004. Occurrence
of mycosporine-like amino acids (MAAs) in the bloomforming cyanobacterium Microcystis aeruginosa. J
Plankton Res 26: 963-966.
MADRONICH S, MCKENZIE RL, BJRN LO,
CALDWELL MM. 1998. Changes in biologically
active ultraviolet radiation reaching the Earths surface.
J Photochem Photobiol B Biol 46:5-19.
MONTERO O, LUBIN LM. 2003. Mycosporine-like
amino acid (MAAs) production by Heterocapsa sp.
(Dinophyceae) in indoor cultures. Biomol Eng 20:
183-189
NAKAMURA H, KOBAYASHI J, HIRATA Y. 1982.
Separation of mycosporine-like amino acids in marine
organisms using reverse-phase high-performance
liquid chromatography. J Chromatog 250:113-118.
NEALE PJ, BANASZAK AT, JARRIEL CR. 1998.
Ultraviolet sunscreens in Gymnodinium sanguineum
(Dinophyceae): mycosporine-like amino acids protect
against inhibition of photosynthesis. J Phycol 34:
928-938.
NEWMAN SJ, DUNLAP WC, NICOL S, RITZ D. 2000.
14
Antarctic krill (Euphasia superba) acquire a UVabsorbing mycosporine-like amino acid from dietary
algae. J Exp Mar Biol Ecol 255: 93-110.
OREN A. 1997. Mycosporine-like amino acids as osmotic
solutes in a community of halophilic cyanobacteria.
Geomicrobiol J 14:231-240.
OSTHOFF SK, WIERMANN R. 1987. Phenols as
integrated compounds of sporopollenin from Pinus
pollen. J Plant Physiol 131:5-15.
PEARL HW, TUCKER J, BLAND PT. 1983. Carotenoid
enhancement and its role in maintaining blue-green
algal (Microcystis aeruginosa) surface blooms. Limnol
Oceanogr 28: 847-857.
PICKETT-HEAPS D, STAEHELIN LA. 1975. The
ultrastructure of Scenedesmus (Chlorophyceae). II. Cell
division and colony formation. J Phycol 11:186-202.
PORTWICH A, GARCIA-PICHEL F. 1999. Ultraviolet
and osmotic stresses induce and regulate the synthesis of
mycosporines in the cyanobacterium Chlorogloeopsis
PCC 6912. Arch Microbiol 172: 187-192.
PROTEAU PJ, GERWICK WH, GARCIA-PICHEL
F, CASTENHOLZ RW. 1993. The structure of
scytonemin, an ultraviolet sunscreen pigment from
the sheaths of cyanobacteria. Experientia 49:825-829.
RIEGGER L, ROBINSON D. 1997. Photoinduction of
UV-absorbing compounds in Antarctic diatoms and
Phaeocystis antarctica. Mar Ecol Progr Ser 160:13-25.
ROZEMA J, BROEKMAN RA, BLOKKER P,
MEIJKAMP BB, DE BAKKER N, VAN DE STAAIJ
J, VAN BEEM A, ARIESE F, KARS SM. 2001.
UV-B absorbance and UV-B absorbing compounds
(para-coumaric acid) in pollen and sporopollenin: the
perspective to track historic UV-B levels. J Photochem
Photobiol B Biol 62: 108-117.
SHAW G, YEADON A. 1964. Chemical studies on the
constitution of some pollen and spore membranes.
Grana Palynol 5:247-252.
SHICK JM. 1993. Solar UV and oxidative stress in algalanimal symbiosis. In: Shima A, Ichihashi M, Fujiwara
Y, Takebe H, editors. Frontiers of Photobiology
Proceedings of the 11th International Congress on
Photobiology. Amsterdam: Elsevier. p. 561-564.
SHICK JM, DUNLAP WC. 2002. Mycosporine-like
amino acids and related gadusols: biosynthesis,
accumulation, and UV-protective functions in aquatic
organisms. Annu Rev Physiol 64:223-262.
SHICK JM, LESSER MP, DUNLAP WC, STOCHAJ
WR, CHALKER BE, WU WON J. 1995. Depth-

Vol. 139 No. 1, June 2010
dependent responses to solar ultraviolet radiation and

oxidative stress in the zooxanthellate coral Acropora
microphthalma. Mar Biol 122:41-51.
SHICK JM, LESSER MP, STOCHAJ WR. 1991.
Ultraviolet radiation and photooxidative stress in
zooxanthellate anthozoa: the sea anemone Phyllodiscus
semoni and the octocoral Clavularia sp. Symbiosis
10:145-173.
SINHA RP, HDER DP. 2002. Life under solar UV radiation
in aquatic organisms. Adv Space Res 30:1547-1566.
SINHA RP, HDER DP. 2008. UV protectants in
cyanobacteria. Plant Sci 174:278-289.
SINHA RP, AMBASHT NK, SINHA JP, HDER
DP. 2003a. Wavelength-dependent induction of
a mycosporine-like amino acid in a rice-field
cyanobacterium, Nostoc commune: role of inhibitors
and salt stress. Photochem Photobiol Sci 2: 171-176.
SINHA RP, AMBASHT NK, SINHA JP, KLISCH M,
HDER DP. 2003b. UV-B induced synthesis of
mycosporine-like amino acids in three strains of
Nodularia (cyanobacteria). J Photochem Phobiol B
Biol 71: 51-58.
SINHA RP, KLISCH M, GRNIGER A, HDER DP.
1998. Ultraviolet-absorbing/screening substances
in cyanobacteria, phytoplankton and macroalgae. J
Photochem Photobiol B Biol 47:83-94.
SINHA RP, KLISCH M, HDER DP. 1999. Induction of
mycosporine-like amino acid (MAA) in the rice-field
cyanobacterium Anabaena sp. by UV irradiation. J
Photochem Photobiol B Biol 52: 59-64.
SINHA RP, KLISCH M, HELBLING EW, HDER D.
2001. Induction of mycosporine-like amino acids
(MAAs) in cyanobacteria by solar ultraviolet-B
radiation. J Photochem Phobiol B Biol 60: 129-135.
SINGH SP, KLISCH M, SINHA RP, HDER DP.
2008a Effects of abiotic stressors on the synthesis
of the mycosporine-like amino acid shinorine in the
cyanobacterium Anabaena variabilis PCC 7937.
Photochem Photobiol 84: 1500-1505.
SINGH SP, SINHA RP, KLISCH M, HDER DP. 2008b.
Mycosporine-like amino acids (MAAs) profile of
a rice-field cyanobacterium Anabaena doliolum as
influenced by PAR and UVR. Planta 229:225-233.
SMITH RC, PREZELIN BB, BAKER KS, BIDIGARE
RR, BOUCHER NP, COLEY T, KARENTZ D,
MACINTYRE S, MATLICK HA, MENZIES D,
ONDRUSEK M, WAN Z, WATERS KJ. 1992. Ozone
depletion: ultraviolet radiation and phytoplankton
biology in Antarctic waters. Science 255:952-959.

SOMMARUGA R, GARCIA-PICHEL F. 1999. UVabsorbing mycosporine-like compounds in planktonic
and benthic organisms from a high-mountain lake. Arch
Hydrobiol 144:255-269.
SOULE T, STOUT V, SWINGLEY WD, MEEKS JC,
GARCIA-PICHEL F. 2007. Molecular genetics
and genomic analysis of scytonemin biosynthesis
in Nostoc punctiforme ATCC 29133. J Bacteriol
189:4465-4472.
STEVENSON CS, CAPPER EA, ROSHAK AK,
MARQUEZ B, EICHMAN C, JACKSON JR,
MATTERN M, GERWICK WH, JACOBS RS,
MARSHALL LA. 2002a. The identification and
characterization of the marine natural product
scytonemin as a novel antiproliferative pharmacophore.
J Pharmacol Exp Ther 303:858-866.
STEVENSON CS, CAPPER EA, ROSHAK AK,
MARQUEZ B, GRACE K, GERWICK WH,
JACOBS RS, MARSHALL LA. 2002b. Scytonemin
a marine natural product inhibitor of kinases key in
hyperproliferative inflammatory diseases. Inflamm
Res 51:112-114.
STOCHAJ WR, DUNLAP WC, SHICK JM. 1994. Two
new UV-absorbing mycosporine-like amino acids
from the sea anemone Anthopleura elegantissima and
the effects of zooxanthellae and spectral irradiance
on chemical composition and content. Mari Biol
118:149-156.
SUH HJ, LEE HW, JUNG J. 2003. Mycosporine glycine
protects biological systems against photodynamic
damage by quenching singlet oxygen with a high
efficiency. Photochem Photobiol 78:109-113.
TAIRA H, AOKI S, YAMANOHA B, TAGUCHI S.
2004. Daily variation in cellular content of UVabsorbing compounds mycosporine-like amino acids
in the marine dinoflagellate Scrippsiella sweeneyae. J
Photochem Photobiol B Biol 75:145-155.
URBACH E, ROBERTSON DL, CHISHOLM SW. 1992.
Multiple evolutionary origins of prochlorophytes
within the cyanobacterial radiation. Nature 355:267270.
VERNET M, WHITEHEAD K. 1996. Release of
ultraviolet-absorbing compounds by the red-tide
dinoflagellate Lingulodinium polyedra. Mar Biol
127:35-44.
VERNET M, NEORI A, HAXO FT. 1989. Spectral
properties and photosynthetic action in red-tide
populations of Prorocentrum micans and Gonyaulax
15

Vol. 139 No. 1, June 2010
polyedra. Mar Biol 103:365-371.

VOLKMANN M, GORBUSHINA AA, KEDAR L,
OREN A. 2006. Structure of euhalothece-362-, a
novel red-shifted mycosporine-like amino acid, from
a halophilic cyanobacterium (Euhalothece sp.). FEMS
Microbiol Lett 258:50-54
VON DER GATHEN P, REX M, HARRIS NRP, LUCIC
D, KNUDSEN BM, BRAATHEN GO, DEBACKER H,
FABIAN R, FAST H, GIL M, KYRA E, MIKKELSEN
IS, RUMMUKAINEN M, STHELIN J, VAROTSOS
C. 1995. Observational evidence for chemical ozone
depletion over the Arctic in winter 1991-1992. Nature
375:131-134.
WHITEHEAD K, HEDGES JI. 2002. Analysis of
mycosporine-like amino acids in plankton by
liquid chromatography electrospray ionization mass
spectrometry. Mar Chem 80:27-39.
16
WHITEHEAD K, HEDGES JI. 2005. Photodegradation

and photosensitization of mycosporine-like amino
acids. J Photochem Photobiol B Biol 80:115-121.
XIONG F, KOMENDA J, KOPECKY J, NEDBAL
L. 1997. Strategies of ultraviolet-B protection in
microscopic algae. Physiol Plant 100:378-388.
XIONG F, KOPECKY J, NEDBAL L. 1999. The
occurrence of UV-B absorbing mycosporine-like
amino acids in freshwater and terrestrial microalgae
(Chlorophyta). Aquatic Bot 63:37-49.
ZHANG L, LI L, WU Q. 2007. Protective effects of
mycosporine-like amino acids of Synechocystis
sp. PCC 6803 and their partial characterization. J
Photochem Phobiol B Biol 86: 240-245.

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