Renewable and Sustainable Energy Reviews 54 (2016) 473481

Contents lists available at ScienceDirect

Renewable and Sustainable Energy Reviews

journal homepage: www.elsevier.com/locate/rser

A review of macroalgae production, with potential applications

in biofuels and bioenergy
Mohsen Ghadiryanfar a, Kurt A. Rosentrater b,n, Alireza Keyhani c, Mahmoud Omid c
a
Department of Mechanical Engineering of Agricultural Machinery and Mechanization, Faculty of Agriculture, Shahid Chamran University, Ahwaz, Iran
b
Department of Agricultural and Biosystems Engineering, Iowa State University, 3327 Elings Hall, Ames, IA 50011, USA
c
Department of Agricultural Machinery Engineering, Faculty of Agricultural Engineering and Technology, University of Tehran, Karaj, Iran

art ic l e i nf o a b s t r a c t

Article history: This review discusses biofuel and bioenergy production from seaweed, and ranges from cultivation to
Received 19 December 2014 nal product, and investigates opportunities, problems, advantages, disadvantages and other issues of
Received in revised form this emerging industry. High levels of structural polysaccharides and low lignin contents make seaweed
21 July 2015
attractive feedstocks for production of liquid biofuels via fermentation and biogas production via
Accepted 19 October 2015
anaerobic digestion. Since macroalgae can be grown in water (oceans and lakes), they will not compete
Available online 11 November 2015
with land-based crops, and thus will not be in competition with human foods. And biofuel and bioenergy
Keywords: production from macroalgae has some environmental benets. Electricity produced from biogas derived
Macroalgae from macroalgae can be cost-competitive to solar thermal, solar photovoltaic and biomass generated
Biofuel
electricity. Biofuel and bioenergy production from macroalgae, however, will entail higher costs than
Bioenergy
terrestrial biomass feedstocks due to higher costs of cultivation and higher costs to remove harmful
Sustainability
Energy security content such as sulfur and nitrogen from the resulting fuel or heavy metals from the residues. Economic
production of biofuels and bioenergy will be available by increasing the scale and efciency of production
of this emerging resource.
& 2015 Elsevier Ltd. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
2. Current macroalgae industry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
3. Chemical compositions of macroalgae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
4. Macroalgae supply chain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475
5. Cultivation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 475
5.1. Hatchery production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476
5.2. On-site growing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476
6. Harvesting methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476
7. Pretreatment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
8. Conversion to biofuel and bioenergy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
8.1. Methane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 478
8.2. Ethanol . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 478
9. Environmental issues. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 479
10. Costs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480
11. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 480

n
Corresponding author.
E-mail address: karosent@iastate.edu (K.A. Rosentrater).

http://dx.doi.org/10.1016/j.rser.2015.10.022
1364-0321/& 2015 Elsevier Ltd. All rights reserved.
474 M. Ghadiryanfar et al. / Renewable and Sustainable Energy Reviews 54 (2016) 473481

1. Introduction 2. Current macroalgae industry

Replacement of fossil fuels by renewable and sustainable
alternative energy has become very necessary due to some pro-
blems such as climate changing, increasing crude oil price, energy
security and limitations of fossil fuel resources [29,14]. Alter-
natives are wind, solar, geothermal, hydroelectric, biomass and
biofuel energy. Currently, the most common types of biofuels are
the so-called "rst generation biofuels", including ethanol, bio-
diesel and pure plant oil (PPO). Sugarcane, corn, soybean, potato,
wheat or sugar beet are the most dominant feedstocks of rst
generation biofuels, some of which may be in competition with
human food resources [5]. Therefore, due to the potential unsus-
tainability of the rst generation fuels, recent efforts have been
focused on the production of "second-generation" or advanced
biofuels. These biofuels are made from lignocellulosic biomass and
agricultural wastes. However, these feedstocks do not have direct
competition with human food resources; but in the case of avail-
able cultivable lands, advanced biofuels production may even-
tually have negative competition with food crops [17]. Algal bio-
fuels are considered third-generation [14]. Since marine crops
like macroalgae could be grown in water, they should not be in
direct competition with land-based foods and crops [47].
In spite of potentially high biomass feedstock amounts of
macroalgae, only a few studies have thoroughly examined this
potential resource [35]. The objective of this study was to review
macroalgae, and to discuss potential applications in bioenergy and
biofuels. This review is focused on biofuel and bioenergy pro-
duction from seaweed cultivation to nal products, and investi-
gates the opportunities, problems, advantages, disadvantages and
other issues encountered related to this emerging industry.
The industrial use of various brown seaweeds has been prac-
tical since the early 20th century, and recently, attentions have
been turned to the production of energy from brown seaweed
resources and also green seaweeds, in particular Ulva spp [7].
In 2008, aquaculture produced aquatic plants at a rate of
approximately 15.8 million tons (by weight) and $7.4 billion USD
in value. Approximately all of these plants were seaweeds (99.6%
by quantity and 99.3% by value in 2008). Major producing coun-
tries are located in east and southeast Asia (in fact, nearly all
producers in 2008 were located here). The most dominant pro-
ducers are China (62.8%), Indonesia (13.7%), Philippines (10.6%),
Republic of Korea (5.9%), Japan (2.9%) and Democratic Peoples
Republic of Korea (2.8%) [18].
The trend of cultivated seaweed has grown in recent decades.
The amount of cultivated seaweed increased from 6.5 million tons
in 2001 to 15.8 million tons in 2010. The value of cultivated sea-
weed increased from US$ 1768 million in 2001 to US$ 4143 in
2010. Two basic types of cultivated seaweed are red (57%) and
brown (43%) [19].
3. Chemical compositions of macroalgae
Chemical composition of macroalgae species is signicantly
different from terrestrial plants. They include lower contents of
carbon, hydrogen, and oxygen and higher contents of nitrogen and
sulfur than that of land-based, lignocellulosic biomass. Addition-
ally, the heating value of various seaweeds is often much lower
than corresponding values for energy crops and other biomass; in
contrast, the ash content is generally higher. Moreover, compared
to terrestrial biomass, they have higher contents of metal and
halogens. Seaweeds and macroalgae often contain bromine and

Table 1
Composition of various seaweeds (macroalgae) and a few lignocellulosic feedstocks found in previous work.a

Components Macrocystisb Laminariac Gracilariad L. digitatae Ascophyllum nodosumf Ulvag Oat Strawh Miscanthush

Water 88.2 88 6782 3.9 1.9

Dry Solids 11.8 12
Proximate Composition, % dry basis
Proteins 17.3 12 11.4 7 2.3 4.63 4.89.8 13.6
Lipids 2 0.53 1.94.8 2.7
Cellulose 5.2 6 3.54.6
Ash 41.1 26 37.7 7 3.6 26.5 1824 30.2 9.4 10.9
Volatile Solids (VS) 58.9 74 62.3 7 3.6 69.8 64.9 74.2
Laminarin 0.8 14 1.26.6
Mannitol 20.2 12 6.810.4
Alginate 15.3 23 2429
Fucoidin 0.2 5 410
Elemental Composition, % dry basis
C 28 34.6 32.4 31.6 42.75 46.32
H 3.92 4.7 4.04 5.22 5.58
O 2.3b 31.2 62.3 38.71 41.79
N 1.86 2.4 0.74 2.18 1.06 0.56
S 1.09 1 0.52 3.1 o .02 o.02
P 0.33 0.35 0.2
K 0.014 0.0096
Energy Content, MJ/kg, dry basis
HHV (higher heating value) 16 g 13.2 17.6g 17.8 19.08
LHV (lower heating value) 11 12.1

a
based upon [44], citing respectively.
b
[9]
c
[41]
d
[40].
e
[48]
f
[3].
g
[4]
h
[47]
 M. Ghadiryanfar et al. / Renewable and Sustainable Energy Reviews 54 (2016) 473481
iodine, whereas land-based biomass often contains substantial
chlorine. They have no lignin and no cellulose [50].
Table 1 shows the composition of macroalgae and a few lig-
nocellulosic feedstocks. Macroalgae has some unique constituents,
including carrageenan, mannitol, agar, laminarin, mannan, ulvan,
fucoidin, and alginate, all of which are not found in either lig-
nocellulosic or microalgae biomass [29]. These compounds have
commercial value and potentially valued long lines [47]. Brown
macroalgae have a great quantity of the carbohydrates laminarin
and mannitol. These two carbohydrates typically occur at about 21
and 26% (dry basis) of Macrocystis and laminaria [47]. The molecule
laminarin (or laminaran) is a -(1-3)-linked-glucan with addi-
tional -(1-6)-linked branches [43]. This molecule is a poly-
saccharide of glucose, serves as a storage molecule. It is abundant in
brown algae [36]. Mannitol is another important component in
seaweeds. It is a sugar alcohol, and is comprised of six carbon
atoms. Compared to most other sugars, mannitol has a lower
caloric value, and has been found to be effective as a sweetener in
various food products, especially for diabetic diets. It has also been
found to have effective pharmaceutical applications, such as
decreasing cellular edema or increasing urinary output [58]. Cell
walls of brown algae are composed of 1025% (dry weight basis) by
the polysaccharide alginic acid (often referred to as algin or algi-
nate). Extracted algin quickly absorbs water (200300 times its own
weight) and is thus effective to use as an additive in dehydrated
products, as well as the paper and textile industries. It has also
found use as a food thickener and stabilizer [3].
Compared to terrestrial biomass, macroalgal contents of water
(7090% fresh wt.) and minerals such as alkali metals (1050% dry
wt.) are high, unlike the low protein (715% dry wt.) and lipid
Table 2
Conceptual supply chain for use of seaweeds to produce biofuels and bioenergy (based in part, upon Bruton et al., 2009).
475
contents (15% dry wt.) [29]. Seaweeds also have higher contents of
sulfur, nitrogen, and other minerals and these contents vary according
to growth stages. Therefore, the optimal time for macroalgae har-
vesting should be selected according to the growth cycle [13,50].
4. Macroalgae supply chain
The supply chain for biofuel and bioenergy production from
macroalgae is summarized in Table 2, based on [7]. The rst step is
biomass generation. It can be collected from natural stocks or
generated via cultivation. Harvesting can be done manually or
mechanically. Biomass pretreatment includes cleaning, desalina-
tion, dewatering and drying (when needed). Depending on the
ultimate use for the macroalgae, downstream processing can vary
[7]. More information about each of these stages follows in the
next several sections.
5. Cultivation
Almost 95% of seaweed used by humans is a result of cultivation
activities [19], and nearly 93% of this cultured seaweed are these four
genera: Porphyra, Undaria, Laminaria, and Gracilaria [61]. Only 6% of
global seaweed comes from natural stocks [7]; the rest is cultivated.
Seaweed production consists of 9 steps, as follows [16], from
which steps 26 are performed in the hatchery, and steps 79 are
done at the grow out phase:
1. Collection of fertile seaweed.
476 M. Ghadiryanfar et al. / Renewable and Sustainable Energy Reviews 54 (2016) 473481

2. Release of spores from fertile material.

3. Development of culture under controlled lab conditions.
4. Induction of reproduction under altered lab conditions.
5. Spraying of fertile cultures on to suitable substrate (culture
string).
6. Development of culture on the string substrate under lab
conditions.
7. Deployment of culture on long line(or other) systems at sea.
8. Development of seaweed in approximately 67 months.
9. Harvest of seaweed.

5.1. Hatchery production

The basic method to cultivate seaweeds nearshore, offshore, or

in ponds is to attach the seaweed seedlings to the lines. These
seedlings require a place attaching them to long lines and also
protecting these young plants before transferring to the main farms.
A hatchery provides protected areas for the seedlings, and facilities
to establish grow out arrays. To minimize the power for pumping
seawater and the transportation cost of seedlings to the sea, the
hatchery must be located on at, low-lying land adjacent to the sea
with a low pumping head. Also, the hatchery must have sufcient
electrical power, road access, space for tanks, a laboratory, an ofce,
and other support facilities [16]. Transition of seaweed cultures to
sea occurs after a laboratory period of 12 months.

5.2. On-site growing

Two general methods for seaweed farming are hanging (ver-

tical) kelp rope system and horizontal kelp rope system (Fig. 1)
[42]. Seaweeds are attached to the horizontal rope or the hanging
rope, which is then linked to a vertical rope that is suspended by
oats and also is attached by an anchor and anchor line to the oor
of the ocean or bay. In the vertical, or hanging, method, the rope
attached to the seaweed is kept vertical by weights attached to its
tip, but in the horizontal system multiple oating lines are con-
nected together by horizontal ropes, to which the seaweeds are
attached. These two systems can be constructed as long line
(single raft unit) or grid (raft block) in commercial seaweed farms
(Fig. 2).
Macroalgal farm cultivation methods include far-shore farms,
in-shore coastal farms, and ponds which is built on land. Currently,
pond culture macroalgae is used for specialty markets, and such
farms are integrated with other aquatic products [47]. Nearshore is
the most dominant method, and so far offshore farming is only in
the experimental phase [7].
Nearshore are the farms in locations near shorelines. Nearshore
farms generally have shallow depth that enables seaweeds to
attach and grow, and also provides a sheltered environment for
aquaculture operations [46]. For example, the most common
method used in Fiji is the single-line or xed off-bottom. The
advantages of this method are: 1. simple to construct, 2. easy to
manage, 3. low cost, and 4. easily accessible at low tide [32].
Offshore farms are those that are constructed in deep water,
requiring growth structures that are anchored to ocean bottoms,
or the oating lines requiring positioning devices. This method is
suitable in protected areas with weak water currents or with too
deep water in which xation the bottom lines is too difcult [54].
Some species of macroalgae such as Sargassum oats on the
ocean surface. This buoyancy allows oating algal farms to be
constructed in the oceans. Thus, the high costs of complicated farm
structures and consequently planting and harvesting costs will be
reduced [9]. Cultivation of the seaweed closer to the water surface
in this method causes faster growth rates but farm managing is
difcult and the seaweed is susceptible to damage and degradation
to weather, waves, and boats [32]. The transportation costs of
Fig. 1. Vertical (a) and horizontal (b) rope systems used for seaweed farming
(based, in part, on Philippsen, 2010).
seedlings and harvested seaweeds to and from the farm, increases
with increasing farm distance from the coast. In terms of the long
lines, ropes and rafts are the major problems of this method. One
suggestion to reduce these problems, and also costs, is to share
infrastructure with a wind farm or other offshore enterprises [7].
Land-based ponds, on the other hand, need to have nutrients
applied. The advantages of this method of cultivation include: 1.
Easier management of plants; 2. Are not always necessary; 3.
Nutrient application can be readily applied; 4. Problems which
occur with open sea cultivation (e.g., weather, waves, etc.); and 5.
These types of farms may be located close to conversion/utilization
operations [9]. However, this method results in much higher
production costs [42].
6. Harvesting methods
Seaweeds can be harvested both manually and mechanically.
Manual harvesting is the most common method globally, and can
be used for both natural and cultivated macroalgae. In manual
harvesting, devices such as sickle, fork, and net are used to uproot
the algae. Mechanized harvesting methods, which can involve
mowing, have been developed. Mechanized harvesters including
rotating blades, suction, or dredging cutters, require boats or ships
to operate [47].
For example, boat with a hook-like gear called a scoubidou is
used to harvest Laminaria digitata in France. This gear turns
around itself and uproots L. digitata. The tool is then rotated in the
reverse direction to release harvested seaweeds inside the boat
[34]. Also specialized boat is applied to harvest Laminaria hyper-
borean. In this boat a large fork-like device drag through seaweed
bed and after nearly two minutes the crane on the boat which is
 M. Ghadiryanfar et al. / Renewable and Sustainable Energy Reviews 54 (2016) 473481
Fig. 2. Both vertical and horizontal systems can be implemented using long line
single raft (a) or grid raft block (b) systems. An example of commercial seaweed
farming is shown in (c) (based, in part, on Philippsen, 2010).
xed to the rake, lifts it and approximately 2 t of seaweed is car-
ried into the boat. Another boat, called a sablier with a suction
dredge, can be used to harvest Lithothamnium calcareum. [34].
The option of using a vertical wet-well was studied by [30].
They attached a vertical wet-well with depth gauges to a hydraulic
arm that moved the wet-well along the bottom of the ocean oor.
The wet-well chopped up the seaweed and brought it up into a net
on to the boat. The machine was kept from cutting to the bottom
of the plant by using depth gauges that kept it 20 cm from the sea
oor. After testing near two different shore lines, they received
very different results: one test with a rate of 0.2 t/h (tonnes/hour),
and another test with a rate of 1.125 t/h. The larger boats and
477
machinery were found hard to bring close to shore and maneuver
to cut the seaweed [30].
There are mechanical harvesters that come pre-made, without
the operator having to install other machinery such as a crane.
Conver, based in the Netherlands, makes equipment that can be
used for seaweed harvesting without changes. The Conver C430H
is a mid-range mowing boat, which is paired with a T-front cutting
tool and push frame could cut seaweed and then push it towards
any net. At a width of 1.5 m, this mowing boat is a small boat and
could maneuver much easier and allow the operator to get nearer
shore, allowing for more seaweed harvested per hour [11].
Alpha Boats has developed two series (SR and FX models) of
boat harvesters. These harvesters have cutting capacity of 1.61
1.83 m depth and 1.832.13 m width. These boats have a cutting
platform like a combine harvester. There are two cutter bars ver-
tically on the lateral sides of platform with another cutter bar
horizontally underneath it. The platform goes down into the water.
The cutter bar cuts the seaweed, and harvested seaweed is picked
up by a conveyer into the middle of platform and is transferred to
the reservoir by boat [2].
7. Pretreatment
Most pretreatment processes occur before substrate extraction
or direct conversion and are as follows [7]:
1. Removing foreign objects (stones, snails, plastic bags, and other
garbage).
After harvesting, seaweed can have some impurities such as
stones, snails and other forms of debris. Removing these foreign
materials is necessary for all subsequent applications, especially
before chopping or milling.
2. Chopping (milling).
Milling (also known as grinding) is used to reduce the particle
size of seaweeds and to increase the surface area to volume of
the particles. Small particles have higher reaction efciencies
(e.g., during anaerobic digestion to produce biogas, fermenta-
tion to produce alcohols, and hydrothermal liquefaction to
produce bio-oils) compared to unmilled macroalgae [47].
3. Dewatering/drying.
Dewatering of wet seaweed to approximately 2030% water
content stabilizes the biomass and allows lower energy con-
sumption during transportation as well as drying. Unlike
microalgae, macroalgae has less need for dewatering. Anaerobic
digestion, fermentation, and hydrothermal liquefaction each
have specic requirements for water. In some cases, fuel-red
boilers are used for drying of seaweed to a powder or meal
form. In such situations, the cost of the overall process increases
greatly because more energy is required to evaporate the water
[7].
8. Conversion to biofuel and bioenergy
Several harvestings of Macrocystis pyrifera in a year, in addition
to rapidly growing to a large size and a greater biochemical
methane potential (BMP) than that of other seaweeds, all make it a
highly suitable potential energy crop among the various seaweeds,
including species such as Laminaria, Gracilaria, Ulva, Macrocystis,
Pyrifera, and Sargassum [41].
Extraction of oils from macroalgae, which can then be pro-
cessed into biodiesel or hydrocarbons, are restricted because of
low lipid contents, so it is expected that biofuel and bioenergy
from macroalgae will be produced via conversion of carbohydrates
[47]. Due to low heating values and high ash and metal contents
478 M. Ghadiryanfar et al. / Renewable and Sustainable Energy Reviews 54 (2016) 473481

(especially alkali metals) of macroalgae compared to the terrestrial
energy crops, conversion technologies that are most tolerant to
ash are likely the most suitable for macroalgae conversion. Con-
sequently, hydrous pyrolysis or anaerobic digestion methods may
be more suitable than anhydrous pyrolysis, gasication, or com-
bustion for the production of fuels or chemicals from these sub-
strates [50].
to lignocellulosic and protein rich waste [51]. The major compo-
nent of gas produced via hydrothermal method are hydrogen,
carbon dioxide and methane, and minor components are ethane,
propane and butane [51]. Table 4 summarizes some results of
hydrothermal gasication of various seaweeds.
8.2. Ethanol

8.1. Methane
Levels of lignin in macroalgae is very low, which makes is
appropriate for the production of biogas via anaerobic digestion
[13]. By increasing the moisture content of macroalgae the cost of
overall process from transport to storage and also energy needed
for conversion can be negatively affected, however by integrating
methane extraction with a Combined Heat and Power (CHP) sys-
tem, the overall efciency can be improved [13]. Methane from this
method is generally more costly than fossil fuels, but it can be cost-
competitive with other sources of energy if rules are enacted to
limit emissions, including carbon taxes, and with subsidies for
biomass energy [10]. Trials demonstrated that biogas production
from seaweed is technically viable, but to commercialize this pro-
cess the raw material cost must be reduced to under than 15% of
current levels [7]. Two species of macroalgae that have high yields
of methane are Ulva (Cladophora and Chaetomorpha) and Macro-
cystis pyrifera, with up to 0.48 and 0.31 m3 CH4 per kg volatile solids
(VS), respectively [22]. Biofuel and bioenergy production from
macroalgae generally results in higher costs than terrestrial biomass
feedstocks for a number of reasons, including high sulfur content.
Also the high nitrogen content may need to be reduced [47]. To use
as a fertilizer, harmful components of seaweed digestate, particu-
larly cadmium, must be removed. Imminodiacetic acid (IDA) poly-
acrylamide cryogel was used by Nkemka and Murto [39] to remove
heavy metals from seaweed leachate before methane production.
Table 3 summarizes results of methane production of some species
by anaerobic digestion method.
Hydrothermal liquefaction (HTL) is another conversion method
suitable for biomasses with high moisture content [55]. HTL
includes using a pressurized reactor with temperatures lower than
400 C to liquefy biomass using some type of catalyst and water.
This method does not require dry feedstocks and also does not
need organic solvents. So energy consumption, which is one of the
major aspects of the conversion process, is reduced [6]. Large
amount of salts in seaweeds can increase the gas yield compared
Over the years, many researchers have examined ethanol pro-
duction from a variety of biological materials. Is has been shown
that many types of macroalgae and seaweed can be used to pro-
duce ethanol. This includes red macroalgae (e.g., Kappaphycus
alvarezii, Gelidium amansii, Gelidium elegans, Gracilaria salicornia),
green macroalgae (e.g., Ulva lactuca, Ulva pertusa), and brown
macroalgae (e.g., Laminaria japonica, Laminaria hyperborea, Sac-
charina latissima, Sargassum fulvellum, Undaria pinnatida, Alaria
crassifolia) [12].
High levels of polysaccharides and sugar alcohols in macroalgae
(such as mannitol and laminarin; particularly in brown algae),
make seaweed very appropriate feedstocks for production of liquid
biofuels, especially via fermentation [47]. Compared with terres-
trial ethanol feedstocks, macroalgae have greater hydrolysable
carbohydrate content. Ethanol production from 1 t of wet seaweed
was estimated to be 29.6 kg by [1], similar to land-based crop
ethanol yield.
The lower content of lignin and cellulose in seaweeds compared
to wood is suggesting that a similar processing approach to cellu-
losic fermentation may not be best suited for seaweed [7]. Because
seaweed's sugars are not simple and easily fermentable, the tradi-
tional fermentation process may not be a suitable conversion
method. The low lipid content reduces the possibility of production
of fatty-acid and biodiesel. So it is required to develop a biochemical
or thermo-mechanical process to break down the polysaccharides
into monomers before fermentation [7]. Saccharomyces cerevisiae
and Zymomonas mobilis are two important microorganisms used for
ethanol production [25]. The glucans in seaweed are easily hydro-
lysable by enzymes because of low or no lignin content of seaweeds
[60]. Laminaran glucose monomers, which are good substrates for
fermentation, can be hydrolyzed by many microorganisms [26];
glucanases are relatively common [3] Table 5.
High amounts of mannitol in algal hydrolysates can be cost-
effective substrates for microbial ethanol production [31]. Instead
of fermentation to generate NADH (Nicotinamide Adenine Dinu-
cleotide), mannitol dehydrogenize enzyme is used to oxidize

Table 3
Biogas and methane production for various seaweeds (macroalgae) found in previous work.

Species Loading rate (g Temperature (C) Retention time Gas production (litre/g VS) By-products Reference
VS/litre/d) (d)
Biogas Methane

L. Digitata 6.84 TPDa 37 15 451.7b (237 KW Electricity 232.6b [11]

and 367 KW heat)
L. Digitata 52 30 221b 133b 7.98 Nc [34]
8.75 Pc
36.6 Kc
Saccharina latissima 37 119 0.223 0.127 [53]
Saccharina Latissimad 37 119 0.268 0.155 [53]
L. hyperborea 1.65 35 24 0.53 0.28 [21]
L. saccharina 1.65 35 24 0.45 0.23 [21]
A. nodosum 1.75 35 24 0.22 0.11 [21]
Macrocystis pyrifera 1.6 35 18 0.2270.31 [20] citing
[9,10]
Ulva Cladophora Chaetomorpha 35 0.35-0.48 [20] citing [22]

a
t/d (dry basis);
b
m3/t dry seaweed;
c
kg/t dry seaweed,
d
steam exploded at 130 for 10 min
 M. Ghadiryanfar et al. / Renewable and Sustainable Energy Reviews 54 (2016) 473481 479

Table 4
Product distributions from hydrothermal gasication of various seaweeds, as discussed in previous work.

Species Processing pressure Gas Generated (g/kg Coke Generated (g /kg Major Compounds Generated in Aqueous Phase (mg/L) Reference
(Pa) seaweed) seaweed)
Phenol Acetic acid Formic acid Glycolic acid

Alaria esculentaa 3  107 315 107 205 134 55 9.7 [48]

Bifurcaria bifurcataa 3  107 504 10 304 23 83 9.3 [48]
Fucus serratus 3  107 367 22 309 45 143 44 [48]
Laminaria digitataa 3  107 342 33 307 72 77 14 [48]

1 h reaction time; 8.3 g sample size; 140 mL water used

a
T 500 C;

Table 5
Estimates of gross global production yields, carbohydrate levels, and potential ethanol production using major terrestrial crops and seaweeds, as discussed in previous work.

Sugarcane Sugar beet Corn Wheat Macroalgae Reference

Estimated world productiona 68,260 47,070 4815 2800 730,000 [3]

DW of hydrolysable carbohydratesb 11,600 8825 3100 1560 40,150 [3]
Potential ethanol productionc 6756 5150 2010 1010 23,400 [3]

a
kg ha  1 year  1.
b
kg ha  1 year  1 (DW denotes dry weight basis).
c
L ha  1 year  1.

Table 6
Ethanol production from various seaweeds (macroalgae) found in previous work.

Species Enzymes and yeast By-product Efciency Reference

Laminaria digitata Celluclast 1.5 L 8.08 kg Na 75 kga [34]

-glucosidase 8.75 kg Pa
Laminarinase 36.6 kg Ka
Saccharomyces cerevisiae 359 kWha electricity
Eucheuma cottonii Saccharomyces cerevisiae 0.33 g/g (65%) [50]
Kappaphycus alvarezii Saccharomyces cerevisiae 1.7 g/L [32]
Laminaria japonica E. coli KO11 0.41 gb [29]
Celluclast 1.5 L
AMG 300 L
Novoprime 959
Viscozyme L
Ulva (Ulva pertusa kjellman) Saccharomyces cerevisiae & Meicelase Glycerol (6.91 g/L) 30 g/L [57]
Alaria (Alaria crassifolia kjellman) Saccharomyces cerevisiae & Meicelase Glycerol (17.2 g/L) 34.4 g/L [57]
Gelidium elegans kuetzing Saccharomyces cerevisiae & Meicelase Glycerol (9.98 g/L) 55 g/L [57]
Laminaria hyperborea Pichia angophorae 0.43 g ethanol (g substrate)  1 [23]
P. angophorae
Kluyveromyces marxinus
Pacchysolen tannophilus
Laminaria hyperborea Zymobacter palmae 0.38 g ethanol (g mannitol)  1 [24]

a
Per ton dry seaweed.
b
Ethanol per g of carbohydrate.

mannitol to fructose, thereupon microorganisms are able to fer-
ment it anaerobically [25]. Table 6 shows the some studies results
in ethanol production from seaweed.
Unfortunately inexpensive enzymes that can break down algi-
nate commercially is not available yet. Also, conventional micro-
organisms cannot ferment alginate effectively [7]. Alginate can be
fermented only by genetically modied microorganisms [42].
Butanol is another energy based product of seaweeds [27,56].
9. Environmental issues
Seaweed harvesting has some environmental advantages and
disadvantages. Depending on the harvesting methods and activities
to access the harvesting sites, these will affect environmental sus-
tainability, including wildlife disturbance or other environmental
impacts [15]. Sustainable harvesting should use machinery and
instruments that leave the vegetative organs for regrowth. Recent
reports have indicated that the depletion of Laminaria digitata in
France may have resulted from different factors including over-
exploitation and increasing seawater temperatures [15]. Over-
harvesting, which is sometimes the result of mechanical harvesting,
can affect the biodiversity of sea ecology. Results of a study that
compared mechanical and traditional hand harvesting methods
[30] did not show substantial differences between environmental
effects of these methods, but in terms of economic efciency and
effectiveness, the manual harvesting was much better than
mechanical harvesting.
Seaweeds can be grown without added nutrition, pesticides,
and also without land utilization [21,54]. Seaweed improves the
water quality in which it is grown. By integrating seaweed and sh
farms, seaweed can oxygenate water while utilizing the ammonia
480 M. Ghadiryanfar et al. / Renewable and Sustainable Energy Reviews 54 (2016) 473481
excreted by sh [38]. Seaweed can also x nitrogen [8,37,48] and
phosphorus [34]. Seaweed farming has socio-economic benets
for coastal communities [54], and is an essential source of
employment and local food production in developing countries. So
using this alternative resource can reduce the pressure on local
sh stocks and other food production.
According to the higher efciency of photosynthesis in mac-
roalgae (6-8%) than in land plants (1.82.2%) [49], and very fast
growth rate of them [13], seaweed has the best potential to
remove CO2. Macroalgae can be a CO2 sink [13,45]. Previous works
has shown approximately 960 kg net CO2 was absorbed during the
cultivation of one ton dry seaweed [37]. Macroalgae cultivation has
other environmental benets too, such as reducing eutrophication,
acidication and global warming potential [8,20,28].
10. Costs
To establish sustainable cultivation, harvest and use of sea-
weeds, the economics as well as the biology are important.
Because of the high labor demand and costly equipment in some
types of seaweed cultivation, the economic return of the product
must always be sufcient to make it worthwhile [16].
In spite of numerous studies in the biology of seaweed, there
are scarce studies about the economics, despite its potential as
food [33] and medical products [52]. The protability of seaweed
cultivation as energy crops have not been conrmed either. But
there are some alternatives to produce seaweeds economically,
such as producing valuable by-products [47] or integrating sea-
weed farms with other aquaculture farms [40]. To have a protable
seaweed farm it is necessary to sell products with higher price (for
example 2/kg wet basis) or to combine with hatchery, producing
a comparatively valuable bivalve such as scallop [57].
In spite of potential of numerous applications of seaweed cul-
ture, it has not taken off globally. One reason is the high cost of
seaweed production [8]. The estimated cost to produce fuel from
macroalgae are currently high. Because production costs are high,
as the industry develops costs will decline as a results of
improvements in efciencies, and as scale increases [47]. Fur-
thermore, the cost of conversion to biofuels or bioenergy must also
be considered. Three of these methods were compared by [47],
and consisted of methane production and subsequent conversion
into gasoline via syngas and methanol; fermentation of the sea-
weed into ethanol, and hydrothermal liquefaction into gasoline or
diesel fuel. Maximum allowable feedstock prices for seaweed was
estimated to be  6, 28, and 6 $ per gallon, respectively, for each
system. Additionally, conversion of seaweeds will potentially
result in valuable long lines such as alginate and agar. The authors
assumed that 15% of the seaweed was used for long lines in the
rst system, and found that the net cost of the methane process
could be reduced by approximately 60%; this resulted in a max-
imum feedstock price of about $12/dry metric ton.
The social and economic dimensions of seaweed farming was
investigated by [54]. Representative systems based on the culti-
vation of K. Alvarezii were selected and compared in Indonesia,
Philippines, Tanzania, India, Solomon Islands, and Mexico. The
results showed that the net return to a four-person family, only in
Mexico and Indonesia, are higher than the International Poverty
Line. The estimated breakeven selling price of electricity produced
from biogas of anaerobic digestion of macroalgae was approxi-
mately 120/MWh ($154 for exchange rate of 1 $1.28) [13]. By
comparing the estimated electricity price with different resources
in 2017 [53], this price is lower than solar thermal and solar
photovoltaic by 251 and 157 $/MWh, respectively, and is compar-
able to biomass generated electricity by 120.2 $/MWh.
Some researchers have examined integrated multi-trophic
aquaculture (IMTA) of various seaweeds, from both economic
and environmental perspectives [8,40,59]. It was discussed by [40]
that by integrating seaweed production with abalone production,
potential ecological impacts may decrease and farm prots can
increase. This study also found potential reductions in nitrogen
levels in adjacent coastal areas, a reduction of greenhouse gas
emissions in these systems, and impacts on natural kelp beds.
11. Conclusions
Macroalgae has great potential for the production of sustain-
able energy; however, the high costs of cultivation and conversion
are obstacles. Increasing the scale and efciency of seaweed pro-
duction, integrating other aquaculture systems, sharing the infra-
structure of other offshore enterprise such as wind farms, reco-
vering and utilizing by-products and coproducts, increasing the
efciency of conversion into biofuel or bioenergy, and also making
policy changes will all affect this emerging source of bioenergy. If
these changes are implemented, the production of bio-based fuels
or energy from macroalgae may be realized.
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