Marine Chemical and Medicine Resources☆

S Ali and C Llewellyn, Plymouth Marine Laboratory, Plymouth, United kingdom

© 2018 Elsevier Inc. All rights reserved.

Introduction 1
Novel Metabolites and Drug Discovery 2
Marine-Derived Nutraceuticals and Food Additives 3
Sunscreens 5
Biocatalysis 5
Bioremediation 6
Microbial Fuel Cells and Biofuels 6
Biomaterials 7
Future Prospects 7
Further Reading 9

Glossary
Actinomycetes Major group of bacteria commonly isolated from low-nutrient environments. Rich source of unusual small
molecules (e.g., antibiotics) and enzymes.
Cyanobacteria Aquatic photosynthetic bacteria widely found throughout nature.
Macroalga(e) Seaweed(s).
Microalga(e) Microscopic single-cell plants.
Nutraceuticals Natural chemicals, usually contained in foods, with potential benefits to human health.

Introduction

The marine environment consists of several defined habitats ranging from the sea surface microlayer which encompasses the
first few microns of the water column, through the bulk water column itself, down to the ocean floor and the subsurface
sediments underneath which can be found hydrothermal vents, cold seeps, hydrocarbon seeps, and saturated brines, as well as a
wide range of mineral and geological variation. It has become increasingly apparent that within all these oceanic layers there is a
diversity of micro- and macroorganisms capable of generating a plethora of previously undescribed molecules through novel
metabolic pathways which could be of value to both industry and the clinic. The biological diversity in some marine ecosystems
may exceed that of the tropical rain forests and this is supported by the presence of 34 out of the 36 phyla of life. This
biodiversity stems from the wide range of environmental conditions to which marine organisms have adapted for survival,
including extremes of pH (acid and alkali), temperature (high and low), salinity, pressure, and chemical toxicity (complex
polycyclic hydrocarbons, heavy metals).
Marine organisms currently being exploited for biotechnology include sponges, tunicates, bryozoans, mollusks, bacteria,
cyanobacteria, macroalgae (seaweeds), and microalgae. These organisms have produced compounds with good activities for a
range of infectious and noninfectious disease with high specificity for the target molecule (usually an enzyme). Targets of marine
natural products which may be clinically relevant include ion channels and G-protein-coupled receptors, protein serine-threonine
kinases, protein tyrosine kinases, phospholipase A 2, microtubule-interfering agents (of which the largest number identified are of
marine origin), and DNA-interactive compounds. In addition to small organic molecules, marine organisms are increasingly being
recognized as a potential source of novel enzymes which could be of industrial and pharmaceutical importance. More than 30,000
diseases have been clinically described, yet less than one-third of these can be treated based on symptoms and only a small number
can be cured. Thus, the potential market for novel marine compounds for clinical development is enormous. In addition to
providing new molecules for direct clinical intervention, the marine environment is also rich in compounds which are finding uses
as natural additives in foods, as nutritional supplements including color additives and antioxidants, and as vitamins, oils, and
cofactors which enhance general well-being. Marine organisms are also increasingly providing new solutions to developments in

☆
Change History: July 2018: S Plummer (Department of Marine Sciences, University of Georgia, Athens, USA) added keywords and updated the Further Reading
section.

Encyclopedia of Ocean Sciences, 3rd Edition https://doi.org/10.1016/B978-0-12-409548-9.11422-8 1

2 Marine Chemical and Medicine Resources

Enzymes
Substrates
Ceramics Biopolymers
Adhesives Energy-rich oils
Nanostructures Microbial batteries
Hydrogen production

Biocatalysis
Materials
science Biofuels

Algal feedstocks
Surfactants
Fluorescent molecules
Heavy metal recovery
Desalination
Pollution detection

Novel
Marine
metabolites
organisms Bioremediation
for
chemistry

Sunscreens
and
Drug cosmetics
discovery

Collagens
Anti-infectives Food
Nutraceuticals Antioxidants
Anticancer additives
Revitalisers
Metabolic diseases

Vitamins Colorants
Antioxidants Texturing agents
Probiotics Essential oils

Fig. 1 Uses for marine organisms and their products in the chemical, pharmaceutical, energy, and environmental industries.

such diverse fields as bioremediation, biocatalysis and chemistry, materials science, nanotechnology, and energy. Some of the
potential uses of marine products are summarized in Fig. 1.
The oceans have long been a source of nutrients, additives, and medicines derived from marine mammals and fish; however, this
article focuses on some of the potential which is harbored in predominantly microscopic organisms which are now being
increasingly studied for novel bioactive compounds and chemicals and may provide a sustainable alternative source for new
compounds and processes.

Novel Metabolites and Drug Discovery

Marine organisms have long been recognized as a source of novel metabolites with applications in human disease therapy.
Particular emphasis has been placed on the invertebrates such as sponges, mollusks, tunicates, and bryozoans, but more recently
advances in genetics and microbial culture have led to a growing interest in cyanobacteria and marine bacteria. For example, a
number of anticancer drugs have been derived from marine sources such as sponges which have proven difficult to cultivate and
their metabolites display a structural complexity which often precludes total chemical synthesis as an option for potential drug
 Marine Chemical and Medicine Resources 3

(A) (B)

Fig. 2 Examples of marine microbial cultures. (A) Bacteria isolated from the English Channel. (B) Microalgal cultures of chlorophytes, cryptophytes, and
haptophytes.

candidates. In recent years, studies have suggested that many of these complex molecules may in fact be the product of microbes
which live in a symbiotic relationship with the sponge and that some of these molecules may be the final product of reactions
carried out by different organisms. A major challenge within marine biotechnology will be to ascertain the nature of the organisms
present in the symbiotic relationship and to identify the pathways involved in metabolite production. A recent advance in molecular
biology with the development of metagenomics has opened up the possibility of organism-independent cultivation of genetic
material and subsequent screening and characterization of that DNA for novel metabolic pathways and enzymes. This provides a
powerful tool for accessing difficult-to-culture microbes which exist in complex symbiotic relationships.
Recent advances in the isolation and culture of marine bacteria using both flow cytometry and microencapsulation-based
methods have yielded a vast array of previously unknown bacteria (Fig. 2). Increasingly, these bacteria are being tested for the
presence of bioactive compounds with activities against a diverse range of human and infectious diseases including cancer, HIV,
hepatitis C, malaria, and those caused by the increasingly drug-resistant common bacterial pathogens (e.g., Staphylococcus aureus,
Enterococcus faecalis, Mycobacterium tuberculosis). For example, the antibiotic abyssomicin C has been isolated from an actinomycete
which was cultured from marine sediment collected in the Sea of Japan. Abyssomicin has been shown to interfere with the synthesis
of the essential cofactor folic acid in bacteria and is active against the methicillin-resistant Staphylococcus aureus (MRSA) pathogen.
Thus, marine bacteria represent a vast untapped source for novel compounds with the potential for development as novel drugs.

Marine-Derived Nutraceuticals and Food Additives

In recent years, there has been an upsurge in the consumption of nutritional supplements such as vitamins and cofactors which are
essential to cellular function. One traditional supplement has been cod-liver oil, which is rich in the omega-3 and omega-6 long-
chain polyunsaturated fatty acids (PUFAs). Fatty acids have long been used as supplements in the aquaculture industry but in recent
years their medicinal value to humans has been proposed. PUFAs have been implicated in enhanced blood circulation and brain
development, particularly docosahexaenoic acid (omega-3), which plays an important role in early brain development and neurite
outgrowth. Although PUFAs have traditionally been extracted from fish oils they do not naturally occur there but rather accumulate
from the diet of the fish. The primary source of PUFAs are marine microbes, and in recent years the isolation and characterization of
microbes which produce these fatty acids have allowed their production in other organisms. At present, considerable effort is being
expended on obtaining high-level production of PUFAs in plants using genes from microalgae with the aim of eventually producing
PUFAs in plants which have traditionally been a source of natural oils (e.g., linseed oil, rapeseed oil). The technology to grow and
extract oils from such plants in large scale already exists and the development of genetically modified plants which can produce
PUFAs in a readily accessible and sustainable form for an increasing market is highly desirable.
4 Marine Chemical and Medicine Resources

(A)

(B) O
OH

HO Astaxanthin
O

OH

HO
Lutein

B,B-carotene
Fig. 3 (A) Any one species of microalgae contains an array of carotenoid pigments, as shown here with the fractionated isolates obtained from the
chromatographic analysis of an Emiliania huxleyi extract. (B) Chemical structures of carotenoids widely used as color additives and antioxidants.

Consumer-led demand for naturally occurring food colorings and antioxidants has resulted in increased interest in photosyn-
thetic pigments and in particular the carotenoids which occur within marine microalgae (Fig. 3). There is, for example, widespread
use of the carotenoid astaxanthin; this is a pigmented antioxidant produced by many microalgae and is responsible for the red color
often associated with crustaceans such as shrimps, crabs, and lobsters. Astaxanthin possesses an unusual antioxidant activity which
has been implicated in a wide range of health benefits such as preventing cardiovascular disease, modulating the immune system as
well as effects in cancer, diabetes, and ocular health. Its antioxidant activities may also have a neuroprotective effect. It has been used
extensively in the feed of farmed fish as a nutritional supplement and is partly responsible for the strong coloration often observed
in farmed salmon, a fish which naturally accumulates astaxanthin in the wild resulting in the pink hue of its flesh. Other carotenoids
 Marine Chemical and Medicine Resources 5

such as beta-carotene and lutein are used widely as food coloring and antioxidants. Inclusion, for example, in the diets of chickens
leads to a darkening of the egg yolk resulting in a rich yellow color.
Another group of pigments of commercial importance is that of the phycobiliproteins; these are used as colorings, in cosmetics,
and as fluorescent dyes for flow cytometry and in immunological assays. More recent research suggests that phycobiliproteins have
anticancer and antiinflammatory properties. A more unusual and unique pigment which is being used increasingly in personal care
products and may also possess anticancer and anti-HIV activity is “marennine,” a blue-green pigment produced by Haslea ostrearia.
Several species of microalgae and, in particular, Haematococcus, Dunaliella, and Spirulina are now grown on large commercial scale to
accommodate the growing demand for natural pigments.
In addition to being rich in phycobiliproteins, Spirulina, a filamentous cyanobacterium, contains a wide variety of nutrients
including potentially beneficial proteins, lipids, vitamins, and antioxidants. Spirulina is also reported to have various beneficial
effects including antiviral activity, immunomodulatory effects, and a role in modulating metabolic function in humans which could
be of value in managing diseases involving lipids and carbohydrates such as diabetes. Furthermore, studies indicate that pretreat-
ment with Spirulina may reduce the toxic side effects observed with some drugs on mammalian organs such as the heart and kidneys.
The marine environment is a rich source for naturally occurring antioxidants and pigments with a diverse range of microorgan-
isms producing a unique and valuable resource. The potential for the discovery of new pigments and other additives which can be
used to replace some of the existing artificial additives currently being used in the food industry is significant.

Sunscreens

The continuous exposure of marine organisms to strong sunlight has resulted in some, primarily the macro- and microalgae,
evolving compounds which provide a very good screen against ultraviolet (UV) light. These organisms have the ability to synthesize
small organic molecules, called mycosporine-like amino acids (MAAs), which are capable of absorbing UV light very efficiently and
thus prevent DNA damage. Over 20 different MAAs occur in nature with a wide range of marine organisms utilizing them, including
corals, anemones, limpets, shrimp, sea urchins, and some vertebrates including fish and fish eggs. MAAs are widely distributed
across the marine environment; however, they can only be synthesized by certain types of bacteria or algae. For example, red
seaweeds and some bloom-forming phytoplankton species are a particularly rich source of MAAs. Studies have revealed that in
addition to their screening ability, some MAAs, such as mycosporine-glycine, have antioxidant properties. The ability of naturally
occurring compounds such as MAAs to act as effective sunscreens has resulted in some interest from the commercial sector as to the
value of these compounds in creams and cosmetics.

Biocatalysis

The ability of enzymes to synthesize complex chiral molecules with high efficiency and precision is of considerable interest within
the pharmaceutical and chemical industries and marine bacteria present a new source for novel enzymes with not only unusual
synthetic properties but also potentially valuable catalytic and structural properties. This arises from the ability of marine bacteria to
grow under extreme conditions such as high and low temperature, high pressure (extreme depth), high salinity, and extremes of pH.
This has opened up the potential to isolate naturally occurring small molecules which are difficult to synthesize in the laboratory
but which could be of value in synthetic organic chemistry as intermediates. The existence of a large number of potentially novel
enzymes in these same organisms which are capable of performing diverse chemical modifications not readily amenable by
standard chemical synthesis also opens the route to novel chemical modification of synthetic molecules using biocatalysis and
biotransformation.
Growth of microbes under these extreme conditions has led to proteins which possess different temperature optima and
improved stability, which has been exploited in the development of new processes and methods such as the polymerase chain
reaction, a method for amplifying specific fragments of DNA, and which depends on a thermostable DNA polymerase isolated from
a thermophilic microorganism. It has been suggested that enzymes which display high salt tolerance may be of value in the
development of enzyme reactions to be performed in organic solvents as they appear to be less prone to denaturing under
dehydrating conditions.
Marine bacteria make up the largest potential single source of novelty in the world’s oceans and of these the major component
are the actinobacteria which includes the actinomycetes. Actinomycetes are readily isolated from the marine environment and
consequently are the best studied of the actinobacteria but the other more difficult to culture members are now being identified
using advanced culturing and molecular techniques. The actinomycetes in particular hold the promise of tremendous diversity and
to date have been underexploited. Terrestrial actinomycetes are responsible for about half of the known bioactive molecules isolated
from natural sources to date and include antibiotics, antitumor compounds, immunosuppressants, and novel enzymes. Conse-
quently, the isolation of new organisms from the environment and their analysis for novel metabolites has been a cornerstone in
drug discovery. In recent years, however, terrestrial organisms have divulged less novelty than before, and advances in microbiology
6 Marine Chemical and Medicine Resources

and genetics have now made the exploitation of marine-derived actinomycetes more attractive. The recognition that the world’s
oceans are rich in biological diversity and that extreme environmental conditions (e.g., high pressure and temperature at deep-sea
hydrothermal vents) have not repressed the development of organisms to form distinct ecological niches suggests that these habitats
will be a rich source of chemical novelty.
Although much emphasis has been placed on isolating organisms from extreme environments in the search for novel
biocatalysts, the general marine environment should not be ignored. Both micro- and macroalgae have been demonstrated to
produce novel enzymes with possible applications in biocatalysis such as the haloperoxidases, enzymes capable of introducing
halogen atoms into metabolites. For example, two species of tropical red macroalgae produce halogenated compounds as a defense
against predators and such compounds are being tested for medical applications. The availability of haloperoxidases with different
catalytic functions would be of use in generating new types of halogenated molecules for the chemical and pharmaceutical
industries.
The realization that viruses are the most abundant biological agents in the marine environment and the discovery of highly
diverse, ancient, giant viruses with genomes comparable in size to the smallest microbes opens up new sources of genetic diversity.
Current indications are that the oceans contain a wide variety of both DNA and RNA viruses with survival strategies which mimic
those of terrestrial viruses yet these marine viruses encode a great many proteins of unknown function. Most marine viruses are
assumed to be bacteriophages because virus particles are most commonly detected in the vicinity of bacteria, and bacteria are the
most abundant organisms in the oceans.
Recent studies have revealed that marine viruses encode unexpected and novel proteins which would not be expected to occur
within a virus genome. For example, the giant algal viruses have been shown to encode novel glycosylases, potassium pumps, and a
pathway for the synthesis of complex sphingolipids. This biochemical diversity indicates that marine viruses could be a rich source
for exploitation in the future for new types of carbohydrate and lipid as well as new proteins and enzymes.

Bioremediation

Pollution of the marine environment is a growing concern particularly with the continuous discharge of both industrial and
domestic waste into rivers and estuaries leading to concerns about the impact such pollution could have on long-term human
health. The discovery of marine microorganisms capable of detoxifying heavy metals and utilizing complex hydrocarbons as an
energy source has provided a new impetus to develop natural solutions to the problems of environmental pollution. However, it
should be remembered that toxic substances are not the only causes of marine distress and that the utilization of fertilizers and the
disposal of sewage can also result in an imbalance in the marine ecology, resulting in the formation of large, often toxic, algal
blooms which although not always a direct threat to human health do lead to widespread ecological damage. Thus, the discovery of
microbes capable of growing in the presence of high concentrations of ammonia could be of value in the treatment of wastewater,
and an understanding of the anaerobic oxidation of ammonia could lead to the development of new chemical processes. The same
organisms also possess unusual metabolic intermediates such as hydrazine and produce unusual lipids which could also be of value
in the search for new chemical intermediates.
Heterocyclic molecules containing sulfur, nitrogen, and oxygen are among the most potent pollutants and inevitably contam-
inate the marine environment to a considerable extent. The use of microbes to degrade and detoxify such compounds is gaining
considerable interest as a process which is environmentally friendly and would represent a long-term solution to removing
heterocyclic contaminants. A particularly rich source of such organisms is the marine environment where growth in close proximity
to sulfur-rich hydrothermal vents or adjacent to hydrocarbon (oil) seeps on the ocean floor has produced a plethora of microor-
ganisms with metabolisms adapted to the utilization of a wide variety of carbon-, nitrogen-, and sulfur-based chemistries. Again,
these organisms are also a very rich source of enzymes with previously unknown characteristics such as unusual substrate specificity,
which could be of great value to the chemicals industry where they could be utilized in the production of new or difficult-to-
synthesize compounds because they can perform reactions which are difficult to duplicate using traditional synthetic chemistry
methods.

Microbial Fuel Cells and Biofuels

The use of marine organisms to produce fuels has also been proposed. The generation of electricity through the degradation of
organic matter has recently been demonstrated to occur in marine sediments and may be mediated by complex communities of
marine microorganisms. These organisms degrade complex organic matter such as carbohydrates and proteins to simpler molecules
such as acetate which are then used by electricity-generating bacteria to reduce metals such as iron and manganese. By replacing the
naturally occurring metals with an anode these bacteria, under anoxic conditions, will supply the electrons needed to produce an
electric current to a cathode linked to the anode by wires and exposed to the oxygen in the water column. It has been suggested that
this type of system could be used to supply the electricity needed to operate equipment in regions where access is difficult and so
 Marine Chemical and Medicine Resources 7

eliminate the need to replace batteries. Microbial fuel cells would be self-sustaining, would not require the preprocessing of fuels to
function efficiently, and would not contribute net CO2 to the atmosphere nor produce toxic waste as with conventional batteries.
Another area of intense study is the development of renewable biofuels with much focus being given to developing terrestrial
plant species to produce the precursors to biodiesel. Microalgae present a potential alternative source of hydrocarbons for the
generation of biofuels as some species naturally produce significantly more oil (per year per unit area of land) than terrestrial oil
seed crops. Several marine species such as Porphyridiuim, Chlorella, and Tetraselmis are currently under investigation as sources rich in
hydrocarbons suitable for biofuel production.

Biomaterials

Another area of interest is the development of novel biomaterials inspired by marine organisms. Areas of particular interest are
the mechanism of calcium- and silica-based structure formation which is found in many phytoplankton, formation of hard
chitinous shells in many larger marine organisms such as oysters and crabs, as well as the very powerful bioadhesives produced
by mussels and barnacles. Proteins form the basis of a number of naturally occurring adhesive molecules which display a
number of attractive features, particularly for clinical and other specialist applications. These features include the ability to
adhere strongly to both smooth and uneven surfaces with a high degree of bonding strength and the ability to form and
maintain bonds in very humid and wet conditions. This bonding ability seems to be strongly linked to the presence of
hydroxylated tyrosine residues (l-dopa; l-3,4-dihydroxyphenylalanine) in such proteins and it is thought that adhesion involves
interactions between the hydroxyl groups and the target surface. The development of powerful adhesives which can cure rapidly
under wet conditions and are nontoxic would be of particular value in the clinic. At present there is considerable interest in using
bioadhesives in the field of ophthalmology where the use of alternatives to sutures in, for example, corneal grafts is desired in
order to reduce the risks of irritation and scarring to the eye following surgery. Another area where the use of bioadhesives is
being actively researched is in drug delivery where the ability to attach naturally occurring polymers which can slowly release a
drug over time would be useful. This is particularly relevant for poorly soluble biological drugs based on antibodies and other
large proteins which can be difficult to administer. The potential contribution that marine-derived biomaterials and bioadhe-
sives could make to such fields is enormous.
Chitin is the second most abundant natural polysaccharide after cellulose and is found in the exoskeletons of crustaceans such as
crabs and shrimp as well as in the cell walls of fungi and cuticles of insects. The deacetylation of chitin produces chitosan, a
biopolymer with great potential in medicine. Chitosan and its derivatives possess numerous applications due to their properties
which include reactive functional groups, gel-forming capability, low toxicity, and high adsorption capacity, as well as complete
biodegradability and antibacterial and antifungal activities. These properties make chitosan particularly attractive in areas of
research such as drug delivery and tissue engineering where a nontoxic, biodegradable scaffold with antimicrobial activity would
be particularly attractive. Both chitin and chitosan can influence the immune system and are being studied extensively as bio-
materials in the development of supports for accelerated wound healing. These chitosan-based materials are also being modified to
improve adhesion to wound sites and for the incorporation of antimicrobial agents to minimize the risk of infection. Chitosan and
its derivatives are also being used to develop scaffolds for applications in tissue engineering to grow cells to form complex structures
which could ultimately be used to replace damaged tissues and organs.
The elaborate silica-based structures (frustules) which are exhibited by many diatoms have been of interest to materials scientists
for many years and recent studies have begun to reveal some of the characteristics that are present in these silica shells, including an
understanding of the proteins and other molecules involved in structure formation (Fig. 4). The highly precise nature of the
structures has led to suggestions that the silica structures can be used directly as either templates for microfabrication or as materials
for use in microprocesses such as filters in microfluidics. By understanding and manipulating the growth environment of any given
diatom it may be possible to modify the precise geometry of the natural silica shells it produces and the resultant frustules could
then be modified using standard microengineering techniques to create new nanostructures with potential applications in the
development of medical devices. The glass-like properties of diatom frustules, the remains of which form diatomite (diatomaceous
earth), have over 300 recorded commercial applications. The fine pores present in the frustules make them especially useful in
filtration processes and the bulk of diatomite is used for this purpose. It has also been suggested that frustules might have
applications in the development of new optical devices.

Future Prospects

The marine environment, representing 70% of the Earth’s surface, is a vast untapped resource for new chemicals and enzymes, often
with characteristics which are considerably different to anything discovered in the terrestrial environment. A major hurdle to the
study and exploitation of this resource has been the inaccessibility of the oceans but advances in science and technology are now
providing new approaches to isolating and characterizing the organisms present. This should lead to a considerable increase in the
number of novel marine-derived chemicals and medicines available to the market in the near future.
8 Marine Chemical and Medicine Resources

(A)

(B)

(C)

Fig. 4 Scanning electron microscope images of (A) the diatom Thalassiosira, (B) the dinoflagellate Gonyualax, and (C) the dinoflagellate Peridinium showing the
intricacies of cell walls as inspirations to novel biomaterials.
 Marine Chemical and Medicine Resources 9

Further Reading
Derikvand P, Llewellyn CA, and Purton S (2017) Cyanobacterial metabolites as a source of sunscreens and moisturizers: A comparison with current synthetic compounds. European
Journal of Phycology 52: 43–56.
Girguis PR, Nielsen ME, and Figueroa I (2010) Harnessing energy from marine productivity using bioelectrochemical systems. Current Opinion in Biotechnology 21: 252–258.
Green DW, Lee J, and Jung H (2015) Marine structural biomaterials in medical biomimicry. Tissue Engineering Part B, Reviews 21: 438–450.
Gullo VP, McAlpine J, Lam KS, Baker D, and Petersen F (2006) Drug discovery from natural products. Journal of Industrial Microbiology and Biotechnology 33: 523–531.
Higuera-Ciapara I, Félix-Valenzuela L, and Goycoolea FM (2006) Astaxanthin: A review of its chemistry and applications. Critical Reviews in Food Science and Nutrition 46: 185–196.
Hussein G, Sankawa U, Goto H, Matsumoto K, and Watanabe H (2006) Astaxanthin, a carotenoid with potential in human health and nutrition. Journal of Natural Products
69: 443–449.
Imhoff JF, Labes A, and Wiese J (2011) Bio-mining the microbial treasures of the ocean: New natural products. Biotechnology Advances 29: 468–482.
Khan Z, Bhadouria P, and Bisen PS (2005) Nutritional and therapeutic potential of Spirulina. Current Pharmaceutical Biotechnology 6: 373–379.
König GM, Kehraus S, Seibert SF, Abdel-Lateff A, and Müller D (2005) Natural products from marine organisms and their associated microbes. Chembiochem 7: 229–238.
Lovley DR (2006) Microbial fuel cells: Novel microbial physiologies and engineering approaches. Current Opinion in Biotechnology 17: 327–332.
Marszalek JR and Lodish HF (2005) Docosahexaenoic acid, fatty acid-interacting proteins, and neuronal function: Breastmilk and fish are good for you. Annual Review of Cell and
Developmental Biology 21: 633–657.
Napier JA and Sayanova O (2005) The production of very-long-chain PUFA biosynthesis in transgenic plants: Towards a sustainable source of fish oils. Proceedings of the Nutritional
Society 64: 387–393.
Newman DJ and Hill RT (2005) New drugs from marine microbes: The tide is turning. Journal of Industrial Microbiology and Biotechnology 33: 539–544.
Shi C, Zhu Y, Ran X, et al. (2006) Therapeutic potential of chitosan and its derivatives in regenerative medicine. Journal of Surgical Research 133: 185–192.
Shick JM and Dunlap WC (2002) Mycosporine-like amino acids and related gadusols: Biosynthesis, accumulation, and UV-protective functions in aquatic organisms. Annual Review of
Physiology 64: 223–262.
Suleria HR, Osborne S, Masci P, and Gobe G (2015) Marine-based nutraceuticals: An innovative trend in the food and supplement industries. Marine Drugs 13: 6336–6351.
Suresh KK, Dahms H, Won E, Lee J, and Shin K (2015) Review: Microalgae—A promising tool for heavy metal remediation. Ecotoxicology and Environmental Safety 113: 329–352.
Suttle CA (2005) Viruses in the sea. Nature 437: 356–361.
Trincone A (2011) Marine biocatalysts: Enzymatic features and applications. Marine Drugs 9: 478–499.
Wei N, Quarterman J, and Jin Y (2013) Review: Marine macroalgae: An untapped resource for producing fuels and chemicals. Trends in Biotechnology 31: 70–77.
Wilt FH (2005) Developmental biology meets materials science: Morphogenesis of biomineralized structures. Developmental Biology 280: 15–25.