Algal Diversity and Commercial
Algal Products
New and valuable products from diverse algae may soon
increase the already large market for algal products
A lgae are an extremely diverse
group of organisms that make
up the lower phylogenetic
echelons of the plant kingdom. A
precise definition of this group is
elusive; they share many obvious
characteristics with higher (land)
plants, whereas their distinguishing
features from other plant groups are
varied and more subtle (Bold and
Wynne 1985). Most of the algae are
photosynthetic (like higher plants)
or are closely related to organisms
that are. Algae perform roughly 50%
of the photosynthesis on this planet
(John 1994) and thus are instru-
mental in supporting the biosphere.
In this article, I review the char-
acteristics and classification of al-
gae and highlight the profound di-
versity of its members.l then describe
some commercially available prod-
ucts derived from algae and a few
product areas in which algae may
make a significant contribution in
the near future.
Diversity of algae
The algae have long been recog-
nized as a heterogeneous group of
organisms, ranging from the micro-
scopic blue-green algae, which are
closely related to Gram-negative
bacteria, to the large, complex kelps,
which can exceed 10 m in length.
Recent molecular biology data sup-
Richard ]. Radmer is president and
founder of Martek Biosciences Corpo-
ration, 6480 Dobbin Rd., Columbia,
MD 21045. © 1996 American Institute
of Biological Sciences.
April 1996
Richard J. Radmer
Production methods
range from
low-technology ocean
farming to cutting-edge
man ufacturing
port the idea that algae are a group
of organisms that have indepen-
dently acquired chloroplasts (intra-
cellular bodies that contain the pho-
tosynthetic machinery; Gibbs 1992).
The taxonomic classification of
algae is still the source of some dis-
agreement; Bold and Wynne (1985)
summarize 14 such classifications,
in addition to their own. Algae are
generally grouped into more than a
dozen major groups, primarily on
the basis of pigment composition,
storage products, and a variety of
ultrastructural features. Table 1 fea-
tures one such classification scheme.
More recently, the techniques of
molecular biology have been used to
determine the affiliations of the vari-
ous algal groups and their relation-
ships to other taxonomic groups.
Some interesting and surprising re-
sults of these studies are shown in
Figure 1. Note that the various algal
groups are scattered all over the
map. The blue-green algae (also
known as cyanobacteria) are prokar-
yotes closely related to many com-
mon bacteria. These algae are also
considered to be the progenitors of
the chloroplasts of some higher al-
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gae and plants (Gibbs 1992), At the
other end of the spectrum, green
algae are closely related to higher
plants. Dinoflagellates and eugle-
no ids have certain characteristics
that are intermediate between
prokaryotes and eukaryotes, and the
term mesokaryotes is sometimes
applied to these algae (Lee 1989).
These mesokaryotes may be more
closely related to the red algae and
the slime molds than to other algal
groups. Although molecular meth-
ods of phylogenetic analysis are new
and controversial, the data to date
indicate that the algae are heteroge-
neous at the molecular level.
The extensive phylogenetic diver-
sity of algae shown in Figure 1 is
reflected in many aspects of their
existence. A few examples include:
Growth mode. Figure 2 illustrates
the multiple means by which vari-
ous algae can derive their metabolic
energy. Although a defining charac-
teristic of algae is their ability to
grow photosynthetically at the ex-
pense of light energy, some algae (a
few percent) are able to thrive in the
dark at the expense of food sources
such as sugar. This so-called het-
erotrophic growth mode is similar
to that used by yeast, fungi, and
bacteria. The use of these growth
modes varies. Some algae can use
only a single growth mode: they are
the obligate phototrophs and obligate
heterotrophs. Other algae are more
versatile: facultative phototrophsl
heterotrophs are able to switch from
one growth mode to the other, al-
though they are generally likely to
263
Table 1. Major groups of algae according to Lee (1989). Three minor groups are structures (as exemplified by the
not tabulated. Nuc = nuclear characteristics (Prokaryote, Mesokaryote, or Eu- kelps). In this article, I divide algae
karyote); Chi = chlorophyll type; PBr = phycobiliproteins (presem in three groups). into two structural groups-the
Data on oils from Lee et a1. (1985). micro algae and the macroalgae.
Microalgae are best seen under a
Characteristics
microscope; the individual organ-
Common name No< Chi PBP Storage product isms are often less than 1 mm in
their largest dimension. Macroalgae,
Blue-green algae Pro , yes Glycogen on the other hand, can be seen with
Red algae Eu a, d yes Glycogen the unaided eye. Less obvious, but
Green algae Eo a, b ou Amylose
Euglenoids Meso a, b ou Paramylon perhaps more fundamental, differ-
Di noflagella tes Meso a, c, 00 Amylose ences in the internal cellular struc-
Cryptophytes Eu a, c1 yes Amylose tures of various algae have also been
Golden algae Eo a, c l ' c, 0O Ch rysolaminari 11, oil described (Lee 1989). For example,
Haptophytes Eo a, (I' (, no Chrysolamirwnn
differences in the organization of
Diatoms Eo <1, (I' (, 00 Chrysolaminarin, oil

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Yellow-green algae Eu a, ( 00 Chrysolaminarin the genetic material reflect the fact
Chloron1onads Eu a, ( no Oil that some algae are prokaryotes
Brown algae Ell a, ( I ' (2 00 Laminarin whereas others are eukaryotes.

Golden algae
Ecological diversity_ The genetic and
Diatoms phenotypic diversity of algae is mani-
EI JKARYOTES Y cHow-green algae fest in their nearly ubiquitous distri-
Chloromonads
Brown algae bution in the biosphere. Algae com-
PLANTS
monly grow in fresh water and

('Jfeen algae
::;.--______-:::~~----------__~;>----~~MES:'Dinoflagellates
OKARY~S seawater, and several species grow
in extremely high-salt environments,
such as the Great Salt Lake, Utah,
ANIMALS FUNGI
and the Dead Sea, in IsraeL Within.
these aqueous habitats, some algae
grow within a few hundred microme-
ters of the water surface, others in-
habit the subsurface water column,
Euglenoids and a few thrive at the limits of the
photic zone (which is often 200-300
m below the surface). Algae also
PROKARYOTES
grow in soils (from rich hum uses to
Plant chloroplasts austere desert sands), inside rocks,

::::~~----------~::::;>--------
Blue-grCt-"Jl algae .....
Plant mitochondria
__J in snow fields, and in more exotic
locations, sLlch as the fur of sloths
and polar bears. Finally, algae can
either be free living or exist in asso-
COMMON BACTERIA
ciation with other organisms, as in
the case of lichens.

ARCHAEOBAcrERIA
Metabolic diversity. Algae produce
many different and unusual bio-
Figure 1. Phylogenetic scheme based on the analysis of ribosomal RNA sequences chemical compoLlnds, including fats,
by Wainright et a1. (1993), modified by Hecht (1993). Common names, rather sugars, pigments, and bioactive com-
than scientific names, are used in an attempt to clarify the relationships for the pounds. The diversity of algal me-
nomaxonomist. Major phylogenetic groups are all capitalized. tabolism is manifest in some un-
usual ways, such as in:
grow better in one of the two modes fixed carbon sources. (See Kaplan et
than the other. Interestingly, some al. [1986] for a more detailed de- • Fish oils. These oils are currently
algae are able to blend these two scription of the various growth being promoted for their beneficial
growth modes, for example, simul- modes of algae.) health effects, primarily on the basis
taneously deriving metabolic energy of their high content of omega-3
from light and cellular building Structural diversity. Algae appear fatty acids. Interestingly, fish can-
blocks from carbon sources such as in nature as single-celled organisms, not efficiently synthesize these fatty
sugars. These hybrid growth modes, as colonies of similar or identical acids and, like humans, must obtain
often referred to as mixotrophy, cells, as filaments (both branched them primarily from their diet. Thus,
provide certain algae with the means and unbranched), as membraneous fish oils are, in reality, algal oils that
to make maximal use of light and thalli, and as complex multicellular have been accumulated through the

264 BioScience Vol. 46 No.4

food chain. Carbon dioxide _ __ ~ _ _ _-Light
• Toxins. Okadaic acid and its de- Water
rivatives cause a human affliction
called diarrheic shellfish poisoning,
which results from the ingestion of Phototrophy
shellfish containing these com- Mixotrophy ALGAE
pounds. These bioactive compounds Heterotrophy

were initially isolated from sponges;
however, subsequent work has dem-
onstrated that the okadaic acid com-
pounds are produced by algae (spe-
cifically, dinoflagellates) closely
associated with the sponges (Shimizu
1993).
Large number of species. Algae rep-
~
Sugars, fals, etc.
Oxygen
Figure 2. Schematic diagram of the grmvth modes used by various algae.
Table 2. Products froIn macroalgac. From .Jensen (1993), except agarose data were
estimated by the author. Phycobiliprotein dara were provided by A. Tsetsis,

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resent a large and virtually unex- Marrek Biosciences Corp., Columbia, Maryland, personal communication.
plored group of organisms. Even
though the phycological community Market value
is small compared with many other Product (million SUS)
scientific disciplines, new species are Nori rood 1800
being identified at a rate of approxi- Wakame Food 600
mately one per week (Radmer and Kombu Food 600
Parker 1994). A recent analysis by Alginates Food products 230
Paper products
John (1994) suggests that the Biomedical applications
roughly 36,000 known species of Carrageenans Food products 100
algae represent only approximately Cosmetics
17% of the species existing today. Pharmaceutical produCfs
Agars rood products 160
These numbers indicate that there Biomedical applications
are currently more than 200,000 Agarose Biotechnology applications more than 50?
algal species worldwide. Seaweed meal Al1lmJI feed 5
Manure ("Macrl") Agriculture 10
Liquid fertilizer Agriculture 5
Algal products Phycobi I iproteins Biomedical uses 2

The modes of production of com-

mercial products from algae range
from, at one extreme, the harvesting
of wild stands of macroalgae with
minimal postha rvest processing, to,
at the other extreme, the intensive
cultivation of microalgae in closed
culture systems (i.e., fermentation
systems or their photosynthetic
equivalent) with extensive post-
harvest processing. Thus, the pro-
duction of algal products spans the
gamut from low-technology ocean
fanning to cutting-edge pharmaceu-
tical-like manufacturing. The major
algal products are currently pro-
duced by what is called the seaweed
industry. This industry is based on
the harvest and use of macroalgae
(i.e., seaweeds), primarily brown
algae, the largest and most conspicu-
ous of the macroalgae, and red al-
gae, a diverse algal group.
In the following sections I de-
scribe a variety of algal products,
grouped on the basis of their pro-
duction mode and product type.
Economic data are also presented,
although their availability is spotty
and their reliability and position
within the distribution chain (e.g.,
wholesale versus retail) varies for
many of the product areas. I also
include a few products and product
ideas that are in development but
not currently available commer-
cially.
Foods from macroalgae. The major
foods derived from macroalgae are
shown in Table 2. The algal biomass
for these products is derived from
wild, managed, or cultivated stands
of macroalgae that undergo a mini-
mal amount of processing after har-
vest. In these cases, the product is
the biomass itself, rather than chemi-
cals extracted from the algae. Any
postharvest processing serves only
to clean and preserve the intrinsic
character of the alga.
Non. The major algal product in
the world today is nori, the algal
blade (called a thallus) of certain
species of the red macroalga Porphyra.
Nori is a primary constituent of
sushi, a Japanese food item that is
becoming increasingly popular in
the West. Because of its history and
commercial success, this product
provides a good example of the
macroalgal production procedures
currently available.
The seaweed industry is often
described as a cottage industry, and
the methods employed in the har-
vesting and drying of the seaweed
are often small scale, traditional,
and primitive (Naylor 1976). In the
case of nori, however, modern tech-
niques introduced in the 1960s have
provided the means to rapidly in-
crease production yields (Oohusa
1993). Nori cultivation is, in real-
ity, a type of farming in which seed-
like propagules, called conchospores,
are seeded onto nori nets, which are
hung in sheltered ocean areas. Be-
fore the mid-1960s, nori cultivation
was limited to shallow, sandy bays,
where the nori nets could be hung
between poles stuck in the bottom.

Apnl 1996 265

After this time, the nets were often
attached to surface buoys, so that
deeper water could be used for cul-
tivation. Around 1970, a system was
developed to raise the nets out of the
water, which allowed the controlled
drying and temporary storage of the
Porphyra thalli. These sophisticated
growth systems and procedures,
coupled with fast-growing cultiva-
tion forms and mechanized methods
for processing, have provided ample
production capacity, and nori sup-
ply has recently exceeded demand
(Oahusa 1993).
Many nori products are avail-
able; of these, toasted nori sheets
are the largest product segment.
With a market value of approxi-
mately $2 billion and a product vol-
ume of 40,000 tons per year (Jensen
1993), nori represents the most suc-
cessful algal product group at the
present time. Although nori is pri-
marily consumed in Japan, Korea,
and China, sales in other countries
are rapidly increasing-US sales in
1991 were estimated to be $20- $25
millioa (Merrill 1993).
Wakame. Another major algal
food product is wakame, a group of
related foods derived from a brown
alga known as Undaria pinnatifida.
This macro algal product has been
cultivated commercially since the
mid-1950s (Yamanaka and Akiyama
1993). Like nori, wakame is pro-
duced mainly in Japan, Korea, and
China, with Korea being the major
producer. Wakame is more exten-
sively processed after harvest than
most other macroalgal biomass
products. The most popular wakame
product is boiled and salted, which
results in the green product most
preferred by consumers.
As with nori, the primary market
for wakame products is Japan, where
it is available in many forms (e.g.,
salted or dried cut) and is used as an
ingredient in soups, salads, noodles,
aad the like. As of 1990, roughly
20,000 tons, with a market value of
$600 million, were sold annually.
Kombu. The third major algal
food product group is kombu, which
is derived from Laminaria japonica
and related species of brown
macroalgae. These algae are col-
lected during the summer, dried
naturally, and then boiled. A vari-
ety of kombu products are produced,
266
which can be served with meat, fish,
or soups or as a vegetable (Lee 1989).
The annual market for these prod-
ucts is approximately $600 million.
Other macroalgal foods. Many
other macroalgae are also used as
human food. For example, the red
macroalga Palmaria paimata, known
as dulse, has been consumed by
shoreline populations of northwest
Europe since approximately the
tenth century (Lahaye and Vigou-
roux 1992). Another dulse, Rhody-
menia sp., is harvested and con-
sumed in parts of North America,
particularly the Maritime Provinces
of Canada (Naylor 1976), where it
is promoted as a sea vegetable.
Naylor (1976) briefly describes ap-
proximately two dozen other macro-
algal foods.
Industrial products from macroalgae:
the Hydrocolloids. Table 2 also sum-
marizes the uses and market values
of the major polysaccharide prod-
ucts derived from algae. These prod-
ucts, also referred to as hydrocol-
loids, make up the major industrial
products derived from algae at the
present time; their combined mar-
ket value is well over 5500 million.
The marine macroalgae from which
these products arc prepared (certain
species of red and brown algae) a,e
harvested from the wild, from man-
aged wild stands, and from culti-
vated beds. Carrageenans and agars
are obtained from different species
of red algae. Alginates are obtained
from species of brown algae. Unlike
the food products described above,
the macroalgal biomass for these
products undergoes extensive extrac-
tion and processing to yield the final
product (Lewis et al. 1988). The
extreme case is agarose, which is
derived from agar (already a pro-
cessed product) by extensive sepa-
ration and purification (Renn 1994).
Alginates. The alginates are salts
of alginic acid; these salts, and the
sodium salts in particular, are also
known as algin. They are polymers
composed of D-mannuronic acid
and L-guluronic acid monomers.
The sequences and proportions of
these constituents vary with the
source of the algin. The major com-
mercial sources of alginates are
brown macro algae, particularly spe-
cies of Laminaria, Macrocystis, and
Ascophyllum. Alginates are typically
recovered from the macroalgal bio-
mass by extracting the insoluble al-
ginic acid salts with hot alkali (so-
dium carbonate). The sodium
alginate is then separated from the
insoluble seaweed residue by filtra-
tion and purified (McHugh 1987).
The primary characteristic of al-
ginates is their ability to form vis-
cous solutions when dissolved in
cold water, which provides thicken-
ing, gel-forming, water-retaining,
and suspending properties to solu-
tions containing them. These fea-
tures underlie their importance in
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food, industrial, and biotechnologi-
cal applications. Approximately
27,000 tons of algi nates, with a value
of $230 million, were sold annually
around 1990 (Jensen 1993).
Carrageenans. The carrageenans
are a complex group of polysaccha-
rides derived from red macroalgae.
Their unifying characteristic is that
they are all made up of galactose-
related monomers (a-1,3-D-galac-
tose and ~-1,4-3,6-anhydro-D-ga­
lactose) to which sulfate groups are
attached. Three major types of car-
rageenans, designated kappa,
lambda, and iota, are used com-
mercially. They arc primarily de-
rived from Eucheuma cottanii,
Chondrus crisp us, and Eucheuma
spinosum. The carrageenans are
typically recovered from the macro-
algal biomass by extraction with
hot water. Subsequent processing
depends on the characteristics of the
product desired (Stanley 1987).
Like the algi nates, the carrageen-
ans can be used to gel, thicken, sus-
pend, and stabilize foods and other
products. Approximately 15,500
tons of carrageenans, with a value
of $100 million, were sold annually
at the beginning of this decade
(Jensea 1993).
Agars. The agars are mixtures of
polysaccharides extracted from cer-
tain red macroalgae. Like the carra-
geenans, their unifying characteris-
tic is that they are all composed of
galactose-related monomers, in this
case, D-galactose and 3,6-anhydro-
L-galactose. The agars also contain
varying amounts of sulfate, pyru-
vate, and methoxy groups, the con-
tent of which vary with the source of
the macroalgal biomass and the sub-
sequent processing procedures. They
BioScience Vol. 46 No.4
are derived primarily from species
of Gracilaria, Gelidium, Ptero-
cladia, Acanthopeltis, andAhnfeltia.
Like the carrageenans, the agars are
usually extracted with hot water.
Subsequent processing steps serve
to generate a concentrated filtrate,
which is allowed to gel; this gel is
then treated, dehydrated, and milled
(Armisen and Galatas 1987).
The ahility of agars to form stable
gels that retain their characteristics
under a range of conditions (e.g.,
temperature, humidity, and chemi-
cal milieu) underlies their value in
many applications. In addition to
their use in foods, agars are the
media of choice to grow and ma-
nipu
ate microorganisms such as
bacteria and yeast. Because of their
special characteristics, agars can
often command a substantially
higher unit price than the alginates
and carrageenans. Annual sales of
these products amounted to approxi-
mately $160 million, on a volume of
11,000 tons, circa 1990 (Jensen
1993).
Agaroses. The agaroses are highly
refined, specialized macroalgal prod-
ucts that have played a pivotal role
in the biotechnological revolution
(Renn 1990). These products are
manufactured by isolating the less
ionic fractions of agar under highly
controlled conditions designed to
minimize lot-to-Iot variation. The
individual products are targeted to
a variety of specialty applications,
each requiring specific quality as-
surance protocols.
The main applications for the
agaroses are in the biotechnology
area, and these products are key
elements in powerful techniques such
as gene mapping (Renn 1990). Be-
cause of the competitive and spe-
cialized nature of this product area,
market data are difficult to obtain;
the value of more than $50 million
per annum in Table 2 is a crude
estimate. The unit value of some of
these products can be impressive,
ranging beyond $25,000/kg.
Other macroalgal products. In
addition to the major products de-
scribed, macroalgae provide several
other products with significant com-
mercial impact. For example, three
agricultural products-seaweed
meal, manure, and liquid fertilizer-
each have annual sales of several
April 1996
millions of dollars annually (see
Table 2). Naylor (1976) describes
some of these products and their
dissemination and use.
An interesting group of high-value
macroalgal products is the ph yco-
biliproteins. These protein-contain-
ing pigments, which are unique to
certain algae, serve as valuable fluo-
rescent tags with many applications
in high-technology areas, such as
flow cytometry, fluorescence-acti-
vated cell sorting, and histochemis-
try (Glazer 1994). The major prod-
uct in this area is R-phycoerythrin,
which is currently derived from spe-
cies of Porphyra, either cultured or
harvested from the wild. Because R-
phycoerythrin exists as part of a
mixture of several different phyco-
biliproteins, sophisticated separa-
tion and purification procedures,
usually involving chromatography,
are required for the production of
the pure product. The high cost of
this process is balanced by the high
value of the product as a biomedical
reagent. The raw material (purified
phycobiliprotein) currently sells for
approximately $5000/g in a mod-
est, $2-million market.
Products from micro algae grown in
open mass culture. The mass cultur-
ing of microalgae dates back to the
1940s. (Shifrin 119841 and Soeder
[1986] provide interesting accounts
of this effort.) Microalgal mass-cul-
turing systems are generally com-
posed of a series of connected shal-
low channels (on the order of 30 cm
deep) called raceways. The algal
suspension contained in the race-
ways is usually mixed by slowly
moving paddles. The management
of these ponds can be complex, and
efficient algal production is at the
mercy of prevailing weather condi-
tions, probably even more so than
macroalgal culture or traditional
agriculture.
The economical harvesting of
microalgae from mass culture ponds
has historically been a problem that
has hindered the commercial devel-
opment of products. Dense cultures
from highly productive ponds gen-
erally contain only approximately
0.1 % of small algal biomass par-
ticles (Shifrin 1984). Even in this
high-yield case, 1000 g of water
must be handled for each gram of
algae harvested, with sophisticated
(and expensive) techniques, such as
flocculation, centrifugation, and
microscreening, used to isolate the
small algal particles. (Compare this
case with the simple harvest of
macroalgae and agricultural prod-
ucts.) Thus, harvesting costs can
contribute substantially to the cost
of the overall process. Successful
microalgal production overcomes
this problem, either by bypassing it
(as in the case of Spirulina) or by
producing a product with a value
high enough to justify these expenses
(e.g., beta-carotene from Dunaliella).
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Because of these pond manage-
ment and harvesting limitations, a
recurring problem in the develop-
ment of microalgal products has been
the high cost associated with cultur-
ing and harvesting microalgae. Al-
though there is likely to be case-by-
case variability, a detailed study by
Borowitzka (1992) indicated that
microalgal biomass costs generally
exceed $10/kg dry weight, exclud-
ing such costs as downstream pro-
cessing. These findings are reflected
in the marketplace; only high-value
products, such as beta-carotene, and
certain algal biomass products that
can command a high price, such as
Spirulina, are viable candidates for
commercialization. Indeed, these are
currently the only commercially suc-
cessful products in this category.
Spirulina. Spirulina is a filamen-
tous blue-green microalga that has a
long history of use in the human diet
(see Richmond 1986). Species of
Spirulina have been consumed as
dried algae cakes since ancient times
in areas such as Lake Chad (in Af-
rica) and Lake Texcoco (in Mexico).
This alga thrives under alkaline con-
ditions and thus can grow as a rela-
tively pure culture, because growth
of most other algae and other or-
ganisms is inhibited. This feature,
coupled with Spirulina's tendency
to float and clump, provides a natu-
rally occurring edible algal bloom
that can be readily harvested with-
out special techniques. Modern
Spirulina production systems make
use of these same features. High-
alkalinity open ponds are inoculated
and maintained to provide condi-
tions for algal growth. The algal
biomass is then harvested and dried.
In the United States, Spirutina is
267
 used primarily as a health food, a
matket niche in which it can com-
mand a substantial price. There is a
good deal of literature, both in peer-
reviewed scientific journals and in
the popular press, concerning the
value and suitability of Spirulina in
the human diet (e.g., Richmond
1986). The safety of Spirulina has
been established through various
toxicological studies (Belay et al.
1993). The annual commercial pro-
duction of food-grade Spirulina was
approximately 800 tons in the early
19905 (Belay ct al. 1993), with reo
tail prices often exceeding $100/kg
(see Table 3).
Beta-carotene. The dominant pig-
ment product currently manufac-
tured from algae is beta-carotene.
This fact is surprising for two rea-
sons: Beta-carotene is a constituent
of all higher plants, and thus is con-
sumed as part of the normal diet,
and it is also produced synthetically
at a significantly lower price. Thus
its viability in the marketplace de-
pends on the consumer's perception
that there are reasons to supplement
the normal diet with this substance
and that there are real differences
between the natural and synthetic
products. Both of these perceptions
are based on data published in peer-
reviewed journals, although firm
conclusions are currently lacking.
Beta-carotene is known to he an
effective antioxidant, which is the
basis for its reputation as a health-
promoting food supplement (Bur-
ton and Ingold 1984). The purported
superiotity of the natural versus syn-
thetic beta-carotene is based, at [east
in parr, on the different isomeric
composition of the two products
(e.g., Makady et al. 1990).
Algal beta-carotene is produced
by growing species of the salt-toler-
ant green alga Dunaliella in ex-
tremely high-salt environments, in
open ponds or raceways, under high
light. These growth conditions pro-
mote the accumulation of beta-caro-
tene to greater than 5% of the total
cell mass. (Glycerol, a low-value
product, is also produced as part of
this process.) The extreme growth
conditions used in this culturing pro-
cess serve to drastically limit the
contamination of the open culture
by other organisms. In fact, these
algal species, along with species of
268
Table 3. Products from microalgac.
Sources are given in the text.
Product Market value
(million SUS)
Spirulin;J Food BO
Bet;J-c;Jrotcnc Nutritional 2S
supplement
Chlorelb Food [00
Labeled Growth media 5 erotrophic algal product is Chia-
compollnds
Spirulina, are the only algae rou-
tinely cultured under open-culture
conditions; in both cases, the ex-
treme growth conditions serve ef-
fectively to exclude other, compet-
1I1g organisms.
The harvest of the carotene-rich
Dunaliella cells from the large, di-
lute culture medium represents a
major effort and expense. The small
algal cells must be separated from
large quantities of high-salt solu-
tion without disrupting the fragile,
wall-less cells. This effort represents
a major technological challenge, and
several clever proprietary techniques
have been developed.
After harvest, the high-carotene
biomass is ptocessed by a variety of
procedures to generate a variety of
products, ranging from crude high-
carotene biomass to high-concen-
tration beta-carotene in a vegetable
oil carrier. Because of the substan-
tial price difference between the
natural and synthetic products
($1400/kg versus $600/kg; see
Radmer and Parker 1994), the algal
products can access only a small
fraction of the $100-million beta-
carotene market (Table 3; Boro-
witzka 1992).
Products from microalgae grown
with fixed carbon. A small percent-
age of algae arc capable of using
fixed carbon, either in addition to
light (mixotrophy) or in total dark-
ness (heterotrophy; see Figure 2).
From a manufacturing perspective,
heterotrophic algal growth provides
major advantages. Fermentation (the
process of growing heterotrophs in
closed systems) is a well-developed
manufacturing technology with a
vast experience base at a variety of
production scales. The cost of pro-
ducing simple products, such as
single-cell protein (microbial bio-
mass used as a food or feed addi-
tive), can be less than $l/kg dry
weight (Crueger and Crueger 1989),
an order of magnitude less than the
cost of phototrophic growth. Equally
important, large fermentation fa-
cilities, with capacities of hundreds
of thousands of liters, are available
virtually worldwide.
Chlorella. The primary het-
rella, a green microalga that has
been produced and consumed in
substantial quantities since the
1960s, primarily in the far East.
Chlorella biomass is produced in
large quantities (hundreds of tons),
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often using a hybrid process in which
the biomass is generated in a closed
fermenter at the expense of glucose
or acetate and is then" greened" by
exposing the algae to light, either in
open ponds or using transparent
plastic tubes. (Soong rJ 9801 gives
an interesting account of this pro-
duction process.) After harvest of
the biomass by centrifugation, the
algal product is dried, using conven-
tional industrial procedures, and put
into its final form.
Chlorella is promoted and sold
primarily as a health food product
in the form of tablets or powder.
Like Spiru[ina, it can command a
substantial price. The annual com-
mercial production of food-grade
Chlorella is at least 1000 tons
(Iwamoto 1993, Soong 1980); with
retail prices often exceeding $1001
kg, the total market value is prob-
ably in excess of $1 00 million (Table
.l ).
Other fermentative products. Sev-
eral other fermenratively produced
algal products are currently in de-
velopment. For example, Running
et al. (1994) have developed a fer-
mentation process for the produc-
tion of L-ascorbic acid (vitamin C)
using a selected strain of Chlorella
pyrenoidosa. This process has not
yet been commercialized.
Heterotrophic algal production
also may playa significant role in
the area of aquaculture feeds. Bi-
valve and shrimp hatcheries require
substantial quantities of algae on a
continuous basis, and the produc-
tion of these algae can be a major
part of the total production costs.
Currently, production costs for pho-
tosynthetic algae used as feeds are
probably greater than $160/kg dry
weight (De Pauw and Persoone
BioScience Vol. 46 No.4
1995). Most of the algal feeds for
these applications are generated and
used in the same commercial entity
(e.g., Fulks and Main 1991). A re-
cent attempt by Celsys (a company
in Cambridge, United Kingdom) to
commercialize this area by supply-
ing heterotrophically produced feeds
was not successful. More recent
work, notably by Gladue and Maxey
(1994), suggests that this area may
still hold promise.
Probably the most exciting recent
development in the algal product
area is the demonstration that valu-
able omega-3 fatty acids, notably
docosahexaenoic acid (DHA), can
be efficiently produced using het-
erotrophically grown algae (Kyle et
al. 1992). DHA plays a vital role in
infant nutrition. It is the predomi-
nant structural fatty acid in the gray
matter of the brain and retina and
must be supplied, preformed, in the
diet, particularly in the case of in-
fants. DBA is present in human milk
but not in most infant formulas cur-
rently on the market. Despite the
need for this microalgal product,
DHA is currently available only in
small (kilogram) quantities (for re-
search) and as a component of some
infant formula products in Europe.
It is not available to consumers else-
where.
Microalgae produced in closed pho-
tosynthetic systems. Many biotech-
nological applications of microalgae
(and probably macroalgae as well)
require the use of closed culture
(Radmer and Parker 1994). For those
cases in which the alga of interest
cannot be grown heterotrophically,
closed photosynthetic systems must
be considered.
The photosynthetic growth of al-
gae in closed systems has been prac-
ticed at the laboratory scale for the
better part of this century, and effi-
cient pilot-scale production units
have been developed as part of the
US space program to supply food
and atmosphere regeneration for
astronauts. (Watson 11979] de-
scribes some of the patent literature
in this area.) However, the use of
these production systems for com-
mercial purposes is in its infancy. A
primary impediment is the high cost
of generating the algal biomass:
Closed systems that make use of
April 1996
sunlight encounter operating costs
at least as high as those endured for
open ponds (Borowitzka 1992),
whereas those using artificial light
arc likely to accrue costs several-
fold higher than the other production
modes (Radmer and Parker 1994).
The only product area that cur-
rently has a high enough value to
offset the high costs of artificially
illuminated closed photosynthetic
algal culture is stable isotope-la-
beled biochemicals. These products,
produced under completely closed
conditions in which the gas phase is
recycled, are finding increasing use
in the areas of protein structure eluci-
dation (particularly as applied to ra-
tional drug design) and noninvasive
diagnostics (Cox et al. 1988). The
major products in this area, uni-
formly labeled I 'C-glucose, growth
media uniformly labeled with 11C
and ])N, and labeled amino acids
and other sugars, sell into a rela-
tively small $S-million market
(Table 3). The unit prices are gener-
ally mnre than $100/g.
Conclusions
The group of organisms collectively
referred to as algae is not a natural
assemblage. Although most algae are
capable of photosynthesis, this ca-
pability was probably acquired by
several diverse taxonomic groups
independently and in various ways.
These polyphyletic origins are re-
flected in the profound diversity of
this group.
Algae currently provide the basis
for a large, multibillion dollar in-
dustry that is largely invisible, par-
ticularly to consumers in the West.
A few notable points are:
• At the present time, food prod-
ucts, sold primarily in the Far East,
comprise the major use of algae when
reckoned on the basis of known
market value.
• The major industrial use of algae
is in the area of hydrocolloids; in
aggregate, this business is a large
one that has a major presence in the
food industry.
• Major components of the biotech-
nological revolution have been based
on the use of agar and agarose, and
the continued development of spe-
cialized agarose has been a vital
aspect in the development of these
technologies.
• Although the algal kingdom is
extraordinarily diverse and may
comprise more than 200,000 spe-
cies, major algal products at present
are based on only 10-20 algal spe-
cies, almost all of them macroalgae.
The development of valuable
products from algae is in its infancy.
Historically, a major impediment in
this endeavor has been the lack of
suitable culture systems, so that the
supply of raw materials was depen-
dent on harvest from the wild. The
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development of culture systems for
the production of macroalgae, pri-
marily in the Far East, has resulted
in a stable supply of many of the
macroalgae of commercial value.
Similar developments in the supply
of microalgae are occurring today.
Algal molecular biology is not
nearly as well developed as that of
yeast or bacteria, and presently there
is no publicly disclosed commercial
development or production system
that uses this technology. Similarly,
the culture of macroalgal tissues as
unicells or multicellular masses is a
topic of current research interest,
but no commercial product is made
in this manner. One might expect
the development of new algal prod-
ucts to accelerate as these technolo-
gies mature. The recent progress in
the cryopreservation and cryo-
storage of microalgae should greatly
aid in the maintenance and perpetu-
ation of high-yielding algal strains.
Although there has never been a
reported case of a commercial
pharmaceutical product discovered
or produced through algae, extracts
of many algae have shown interest-
ing biological activities in a variety
of bioassays, and many interesting
and novel chemical entities ha ve been
isolated and characterized during
the course of this work. Product
development in this area, as in many
others, has been hindered by our
inability ro culture the algae of in-
terest under controlled conditions
at a large scale. Progress in this area,
coupled with the ever-growing tools
of biotechnology, should facilitate
the production of additional useful
and valuable products from this di-
verse and fascinating group of or-
gal11sms.
269
Acknowledgments
I thank Bruce Parker and Teri
Watson for their help in preparing
this article.
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