Renewable and Sustainable Energy Reviews xx (xxxx) xxxx–xxxx

Contents lists available at ScienceDirect

Renewable and Sustainable Energy Reviews

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

Offshore macroalgae biomass for bioenergy production: Environmental

aspects, technological achievements and challenges
Francois Fernanda,b, Alvaro Israelc, Jorunn Skjermod, Thomas Wicharde, Klaas R. Timmermansf,
⁎
Alexander Golberga,
a
Tel Aviv University, Porter School of Environmental Studies, Tel Aviv, Israel
b
Aix Marseille University, Department of Physics, Marseille, France
c
Israel Oceanographic & $2 Limnological Research, Ltd., Haifa, Israel
d
SINTEF Fisheries and Aquaculture, Trondheim, Norway
e
Friedrich Schiller University Jena, Institute for Inorganic and Analytical Chemistry, Jena, Germany
f
NIOZ Royal Netherlands Institute for Sea Research, Texel, Netherlands

A R T I C L E I N F O A BS T RAC T

Keywords: Economic and environmental developments in the last decades call for the displacement of fossil fuels to
Algae biofuel alternative energy sources. Biofuels are considered as a part of the solution for this challenge. Sustainable raw
Bioenergy materials for the production of transportation biofuels such as biodiesel, biobutanol and bioethanol, can be
Biorefinery obtained from algal biomass. In particular, marine macroalgal biomass is a promising feedstock for
Macroalgae
transportation biofuels because of (the)its fast growth and its potential cultivation on seawater, avoiding
Offshore
Renewable energy
competition of resources with conventional agriculture of terrestrial plants used for food. In addition, dissolved
Biofuels inorganic nutrients like nitrogen, phosphorous and carbon are taken up by macroalgae, helping to alleviate
Fermentation eutrophication in seas and oceans. Using biological, chemical and engineering advances in the last decades,
Aquaculture technologies to provide cost efficient cultivation, harvesting, extraction and processing of sustainable biofuels
Policy making have to be elaborated. This paper provides a review of macroalgae based biorefineries with offshore cultivation
Seaweed cultivation and consequent biomass conversion into transportation liquid biofuels. We discuss the methods for offshore
cultivation, harvesting, and conversion of macroalgae biomass into liquid transportation biofuels. Based on the
current information and global experience, we present local perspectives specific for France, Germany, Norway,
the Netherlands and Israel on the potential use of Exclusive Economic Zone for transportation biofuels
production. Marketable suggestions for future research need to include all stakeholders of a given site for
offshore biorefinery development.

1. Introduction of derived chemicals and bioenergy. Design of a sustainable biorefinery,

Sustainable production of food and generation of energy are the
major challenges for the world for the next decades [1]. One of the
approaches for these challenges is to increase the use and efficiency of
the solar energy. Therefore, economically efficient, socially and envir-
onmentally sustainable conversion of solar energy into valuable
products is a major contemporary challenge for science, governments
and businesses worldwide. Transportation fuels, electricity, heating,
cooling, drinking water, food, animal feed, chemicals, and materials are
all potential products of solar energy conversion. Biorefining is one of
the pathways to convert solar energy into these useful products.
Biorefinery integrates the capture of solar energy and carbon dioxide
(via photosynthesis), biomass harvesting, processing, and distribution
which will generate sustainable food, fuels and chemicals is a complex
task and is largely influenced by local raw material supplies, advances
in multiple technologies and socio-economic conditions [2].
The choice of raw biomass material is critical to ensure the efficient
production of biofuels [1]. The currently used crops and cultivation
methods supplied raw biomass for the food and feed sectors for
hundreds of years; however, most recently they also started to supply
biomass for the production of transportation fuels. First generation
liquid biofuel feedstocks includes traditional agriculture crops: cereals,
potatoes, sugar beet, rapeseed, wood and dedicated energy crops; while
first generation fuel products include ethanol and biodiesel [3]. The
second generation biomass feedstocks include animal fat, and dedi-
cated lignocellulosic crops; and produce hydrotreated vegetable oil,

⁎
Corresponding author.
E-mail address: agolberg@tauex.tau.ac.il (A. Golberg).

http://dx.doi.org/10.1016/j.rser.2016.10.046
Received 18 January 2016; Received in revised form 25 October 2016; Accepted 26 October 2016
Available online xxxx
1364-0321/ © 2016 Elsevier Ltd. All rights reserved.

Please cite this article as: Fernand, F., Renewable and Sustainable Energy Reviews (2016), http://dx.doi.org/10.1016/j.rser.2016.10.046
F. Fernand et al. Renewable and Sustainable Energy Reviews xx (xxxx) xxxx–xxxx

cellulosic-ethanol, biomass-to-liquids (BtL)-diesel, bio-butanol, and

advanced drop-in replacement fuels such as fatty-acid ethyl esters,
alkanes, alkenes, terpenes and methyl ketones [4–10]. Recent studies,
however, indicate the future of biomass sector development is under
high degree of uncertainty mainly due to the limited crops yields and
land availability [1,11]. With increasing world population, arable lands
will be in full intended to food cultivation. Producing biofuel from the
farming of traditional food crops appears awkward regarding food
security and freshwater consumption [12,13].
Alternative biomass supply can come from micro- and macroalgae.
Because of their remarkably high growth rates compared to most other
photosynthetic organisms, the use of marginal land for cultivation,
valuable commercial cellular compounds and potential high lipid
accumulation, microalgae have been the focus of intense research in
the last 50 years [14]. However, cost-effective cultivation, harvesting
and dehydration difficulties currently still prevent broad scale, sustain-
able microalgae technologies implementation for biofuel production Fig. 1. Conceptual offshore biorefinery system for chemicals, food and energy produc-
[14]. tion. Image adapted with permission from Global potential of offshore and shallow
waters macroalgal biorefineries to provide for food, chemicals and energy: feasibility and
Marine macroalgae, also ranked among the most efficient photo-
sustainability Lehahn Y, Ingle, K.N, Golberg A. Algal Research. 17:150-160,2016.
synthetic organisms on earth, bear valuable chemical compounds.
Macroalgae must be attached and are, thus, restricted to the seafloor
2. Environmental conditions that predicate macroalgae
(benthos) or other substrate within the photic zone. In addition to
biorefineries efficiency
photons to activate the photosynthetic machinery, macroalgae also
need nutrients such as nitrogen, phosphorus as well as inorganic
Multiple environmental parameters control successful macroalgae
carbon to grow. In order for these nutrients to be available and enable
cultivation. Water quality, photosynthetic active radiation (PAR),
algal growth, they need to be dissolved and present in enough
temperature, salinity, concentrations of inorganic nutrients including
concentrations in the seawater medium. Nitrogen can be found in the
CO2, and absence of environmental toxins are required for effective
sediment or in seawater as nitrate or ammonium [15], the latest being
biomass accumulation via photosynthesis [30–32]. These environmen-
the preferred nitrogen source. Dissolved inorganic phosphorus enters
tal parameters are dynamic and depend on natural and human factors.
seas and oceans from terrestrial sources. The source of inorganic
For example, we can expect temperature and light to vary throughout
carbon is air-born CO2 that dissolves into seawater, and due to the
the day, and during seasons, together with other equally important
generally high seawater pH, creates a significant pool of available
events such as wind, rainfalls and tides, which will all affect the
HCO3- for algal photosynthesis. In addition, certain macroalgae can
macroalgal growth [33]. Humans also play a role through landform
take up atmospheric CO2 as well. Overall, these features are of major
nutrient enrichment [33,34], aquaculture [35] and wastewaters [36].
interest regarding two points. First, since marine macroalgae fix
Variable environmental conditions also impact the final product since
inorganic carbon, they can be used as carbon trap and then as fuel.
the chemical composition of the macroalgal tissues will strongly be
Second, high levels of nutrients dissolved in seawater (primarily in
affected by the prevailing growth environment [37]. More recently,
coastal eutrophic waters) can be recycled through algal growth and
global changes occurring in the marine environment are forcing the
subsequent harvesting of the biomass yields. Indeed, an expanding
geographical redistribution of macroalgae on a global scale, for
body of evidence has demonstrated that marine macroalgae, which
example the retrieval of kelps from warming waters within the North
contain very little lignin and do not compete with food crops for arable
Atlantic shores [38], challenging the prediction and planning of the
land [16] or freshwater [17], can provide a sustainable alternative
high-yield areas.
source of biomass for the production of food, fuel and chemicals, such
as bio-ethanol and bio-butanol [14,18–24].
Producing sustainable biomass offshore for bioenergy is promising 3. Species choice, cultivation and harvesting methods
because of its sustainability, but extremely challenging endeavor.
Although the concepts of Ocean Farms (Fig. 1) have been introduced As benthic organisms macroalgae are typically attached to hard
decades ago, current macroalgal cultivation is mostly suitable and substrates, yet a few species can also grow floating in the upper layers
therefore practiced in protected, near shore areas. The current concepts of the seawater surface [53]. Naturally growing macroalgae are
of offshore marine biomass cultivation include near farm concepts for harvested in the subtidal zone or at shallow water, and the harvesting
kelp growth [25], tidal flat farms, floating cultivation [25], ring methods depend on the species. Several techniques exist ranging from
cultivation [26] and most recently wind-farm integrated systems rudimentary ones like hand-picking or cutting subtidal thalli to
[27]. Currently, there is no universal definition for offshore cultivation. bulldoze, tractor or boat harvesting [54]. Skimmer boats have been
However, in most cases offshore cultivation means the movement of developed to harvest macroalgae easily far from the coast [20,55], such
farm installations from near shore, sheltered environments and facil- as for Laminaria digitata (harvested by a boat with a gear called
ities to more exposed environments, where then frequent harvests “Scoubidou” developed in 1960´s and used by e.g. CEVA – Centre
could have additional logistical and cost implications [28,29]. d’Etude et de Valorisation des Algues) (Fig. 2). Harvesting naturally
The goal of this article is to provide a comprehensive overview on growing macroalgae may have severe environmental impacts unless a
the available technologies and methods of offshore macroalgal cultiva- certain harvesting cycle depending on the macroalgae species is
tion, and the subsequent conversion of their biomass into transportable followed to allow recovery [45,56]. Several methods have been used
biofuels. Hence, this article presents for the first time a comprehensive to increase macroalgae growth rate and to ease the harvesting process.
state of the art of offshore bioenergy production from macroalgae, Table 1 presents the biomass yields for several types of macroalgae
hoping to provide a solid baseline for future developments aiming at achieved with various offshore cultivation methods.
exploring this “old” and “new” source for bioenergy production. There Attaching seaweeds to ropes, lines or nets is a popular way of
is a strong interest by stakeholders, public and policy in a trusted cultivation since installation and maintenance costs are very low. The
sustainable infrastructure providing renewable green energy. cultivation can be done by attaching the seedlings directly to the ropes

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F. Fernand et al. Renewable and Sustainable Energy Reviews xx (xxxx) xxxx–xxxx

(Fig. 3a) [57]. Another approach is via transplantation: seedlings are
grown indoors, then cultured in greenhouse tanks and finally the small
fronds are transplanted onto ropes in the sea [59]. By controlling the
life cycle under laboratory conditions defined feedstock for certain
macroalgae can already be prepared [67,68]. The ropes should be
tensed to reduce diffusion boundary layers surrounding the thalli, thus
increasing water exchange for efficient nutrient availability and fast
growth [69,70]. Horizontal and vertical ropes systems were also
discussed by [57]. A system with concentrically horizontal ropes was
tested in [26] (Fig. 3b). Seabed planting can be cited as well as cage
system (open-topped cage anchored to the sea floor) for near shore
Fig. 2. Boat harvesting of Laminaria hyperborea outside the coast of Mid-Norway
(Photo: Institute of Marine Research, Norway).
cultivation [64]. Co-management with other offshore systems like wind
farms for instance was considered [71–74] and this would be advanta-
geous environmentally and economically [75]. However, there are still
ongoing discussions how extensive open ocean aquaculture can be
developed as multi-use installations within wind farms relying on the
sufficient co-management of different stakeholders [71] (Fig. 3c).
A critical decision in the offshore biomass production for biofuels is
the species choice. Before the appropriate algal species and cultivation
site are selected, a complex interaction of physical, chemical and
biological factors have to be considered. Most importantly, the
intended application must be clear and well defined beforehand. For
example, different macroalgal species could be chosen for their
production of low-cost fuel in combination with “high value” com-
pounds and/or bioremediation applications, where an excess of
nutrients can be converted in biomass for harvest and economic goods.
Important constrains include light, temperature, nutrients, current
velocity but also the capability to resist the harsh conditions such as
high waves and extreme currents in offshore waters. Nitrogen has often
been indicated as the primary limiting factor for seaweed growth,
however phosphorus may also limit production in some systems
(Fig. 4) [76].
Although red algal species (for example Gelidiales), and in a less
extent also green algal species such as Ulva can continue growing
under nitrogen depleted conditions upon extensive uptake of nutrients,
seasonal variability in the ambient nitrate concentration of seawater
will affect the growth of Laminaria species [77]. Furthermore, the life
cycle of a macroalga as well as its mutualistic and competitive

Table 1
Biomass yields of seaweeds cultivated offshore as reported from 1980 to 2015.

Cultivation system Species Location Yield Units Reference

Rope, vertical Undaria pinnatifida North Western coastal bay of 8.3 kg ww/m/139 d [57]
Spain
21a t ww/ha/year [57]

Rope, horizontal Undaria pinnatifida North Western coastal bay of 5.9 kg ww/m/147 d [57]
Spain
Rope (concentrical) Laminaria saccharina German North Sea 4 kg ww/m/6 months [26]
Ropes farm, horizontal Laminaria japonica Hokkaido, Japan 106 t ww/41.2 km2/year [58]
Rope, horizontal, transplantedb Saccharina latissima Northern Spain, Bay of Biscay 7.8 kg ww/m/106 d [59]
45.6 t ww/ha/106 d [59]

Rope, horizontal Laminaria saccharina British Columbia, Canada 3–8 kg ww/m/8 months [60]
Rope, horizontal Laminaria groenlandica British Columbia, Canada 2.6–20.5 kg ww/m/18 months [60]
Rope, horizontal Cymathere triplicata British Columbia, Canada 1.1–2.7 kg ww/m/7 months [60]
Rope, verticalc Palmaria palmata Northwest Scotland 1 kg ww/horizontal meter of top [61]
rope/year
Rope, verticalc Saccharina latissima Northwest Scotland 28d kg ww/horizontal meter of top [61]
rope/year
Rope, horizontal Alaria esculentae Ard Bay, Carna, Co. Galway, 45.7 kg ww/m/year [62]
Ireland
Rope, horizontal Saccharina latissima Isle of Man, Irish Sea 2.8 kg dw/m/year [63]
Cagef Gracilaria tikvahiae Indian river lagoon, Florida 9.7g g dw/m2/day [64]
Cagef Gracilaria tikvahiae Hutchinson Island, Florida 22.4 g dw/m2/day [64]
Nylon line attached to stakes fixed in sea Eucheuma spinosum (Bohol) Zanzibar Island, Tanzania 5.4–7%h daily growth rate [65]
bottom
Floating raft with rope Sargassum naozhouense (Tseng Liusha Bay, Xuwen, Guangdong, 1750 kg ww/km/95 d [66]
et Lu) China
i
Rope net with two bamboo poles Ulva prolifera Jiangsu coastline, China 198.6 kg ww/ha/5 months [41]
Rope net with two bamboo poles Ulva intestinalis Jiangsu coastline, China 89.2i kg ww/ha/5 months [41]

ww: wet weight; dw: dry weight

a
Estimated value.
b
Transplants were 2.1 kg fresh wt m-1 rope.
c
Droppers 1 m apart, with one 10 cm section of seeded string for every 1 m of dropper to 7 m depth.
d
Highest mean yield obtained for a longline.
e
High yielding strain.
f
2 cm plastic mesh on 2.5–5.0 cm diameter PVC pipe frames measuring 1 m x 1 m x 0.25 m deep (0.6 m2 cage).
g
Average between two stations (7.8 and 11.6 g dw/m2/day).
h
Minimum and maximum of five test plants at different locations.
i
Average of six stations.

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F. Fernand et al. Renewable and Sustainable Energy Reviews xx (xxxx) xxxx–xxxx

Fig. 3. Current strategies for offshore biomass cultivation. (a) Harvesting from Seaweed Energy Solutions AS' pilot seaweed farm of 300×300 m in Norway, where two cultivation
systems for kelp are tested: (1) A submerged 200 m long carrier line with a single, continuous cultivation rope for vertical cultivation (front), (2) A two-dimensional carrier (back) for
horizontal cultivation (Photo: SINTEF Fisheries and Aquaculture). (b) The offshore-ring design. The successful ring design for the culture of Laminaria at offshore locations. The major
elements of the system design are magnified: (A) central guide ring with attached carrier rope and culture line, (B) the transition between central steel cable of the mooring and that of
the lower crow’s foot, (C) metal cuffs, to which the crow’s feet and the carrier ropes are attached. From [26] with permission. (c) Multipurpose offshore system concept including, for
example, the Submersible Offshore Shellfish and Seaweed Cage (SOSSEC)-system is shown. With permit from © Bela H. Buck, AWI, Alfred-Wegener-Institut, Bremerhaven (Germany).

increasing biomass abundance the availability of both light and

Fig. 4. Concentrations of dissolved nitrate (µmol L–1) in winter, originating from the
rivers Rhine and Meuse along the coast of the Netherlands, calculated using the ERSEM
model [130].
interactions with other organisms should be investigated in laboratory
studies and proven later on in field studies. For instance, spontaneous
sporulation (generally triggered by sudden changes in ambient sea-
water medium [68]), or the switch from vegetative to sexual life cycle
and the subsequent release of germ cells may jeopardize massive
offshore biomass production.
As the offshore cultivation site should provide non-limiting growth
conditions in the first place, as discussed above, macroalgae growth
depends on saturation kinetics by light intensity, ambient dissolved
inorganic nutrient concentrations and temperature [78]. In this context
it is noteworthy that the biomass of polycultures often increases
compared to monocultures as diversity might enhance cover and
stability of wild macroalgae assemblages [79]. However, with an
nutrients decrease sharply, subsequently inducing decreasing growth
rates [29]. In addition, an associated microbiota, as shown for green
tides, will be affected [80]. This in turn could be critical for long-term
stability of culture, as bacteria can support macroalgae growth,
development and morphogenesis by waterborne bacterial compounds
[68,81,82]. Finally, fouling of the cultivated macroalgae by macro-
scopic organisms like bryozoans, epiphytic macroalgae, hydroids, snails
and blue mussels as illustrated for the sugar kelp Saccharina latissima
in late spring and during summer, induce deterioration of the algal
tissue and cause great biomass loss [83]. The impact of such biofouling
is that the biomass has to be harvested at late spring or early summer,
thus limiting the cultivation period and the possibilities for accumula-
tion of storage polysaccharides during the summer months. For
production of a carbohydrate rich kelp biomass aimed for bioenergy
purpose cultivation strategies that reduces or bypasses biofouling is
crucial. In any case as nutrient levels drop, the importance of water
turbulence and wind velocity around the seaweed becomes important
for the acquisition and uptake [84]. Frequent harvesting and thinning
the biomass density can solve the growth limitation also due to limited
light penetration, but will cause additional logistical and cost implica-
tions for offshore facilities [29]. In addition moderate water velocity
substantially improves macroalgal production by increasing the avail-
ability and algal uptake of nutrients including inorganic carbon [70].
4. Technologies for macroalgal carbohydrates fermentation
and bioenergy conversion
While biomass rich in lipids will be suitable for the production of
biodiesel, bioethanol can be obtained more easily from sugar rich
sources. In this part, Tables 2–4 list the different yields obtained for
ethanol, butanol and biodiesel conversion respectively, the type of
macroalgae and the process. We can note that the low fraction of lignin
in macroalgae ease the degradation and for that reason pretreatments
do not need to be as harsh and energy consuming as for more resistant

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F. Fernand et al. Renewable and Sustainable Energy Reviews xx (xxxx) xxxx–xxxx

Table 2
Macroalgae conversion into ethanol: pretreatment, hydrolysis and fermentation processes (reports from 2000 till 2015).

Algal Species Mode Hydrolysis/pretreatment Fermentation Yield (ethanol) Reference

Eucheuma cottonii SHF 5% H2SO4, 100 °C, 2 ha Saccharomyces cereviceae, 28–30 °C, 4.6% [105]
5–6 days
Kappaphycus alvarezii SHF 2.5% H2SO4, 100 °C, 1 h Saccharomyces cerevisiae (NCIM 2.46% [106]
3523), 96 h
Kappaphycus alvarezii SHF 0.2 M H2SO4, 130 °C, 15 min Saccharomyces cerevisiae, 24 h 1.1–1.9 g/L [107]
Gracilaria salicornia1 SHF 2% H2SO4, 120 °C, 30 min. Cellulase, 40 °Cb Escherichia coli (KO11), 30 °C, 30 h 79.1g/1 kg [108]
Gelidium elegans SHF 2% H2SO4, 121 °C, 30 min. Meicelase, 50 °C, 120 hb Saccharomyces cerevisiae (IAM 55 g/L [109]
4178), 30 °C, 48 h
Ulva pertusa SHF Meicelase, 50 °C, 120 h (twice)c Saccharomyces cerevisiae (IAM 30 g/L [109]
4178), 30 °C, 24 h
Alaria crassifolia SHF Meicelase, 50 °C, 120 h (twice)c Saccharomyces cerevisiae (IAM 34.4 g/L [109]
4178), 30 °C, 24 h
Laminaria hyperborea SHF Water pH 2, 65 °C, 1 h. pH 4.5, 121 °C, 20 min Pichia angophorae, 30 °C, 75 h, 0.43 g/g substrate [110]
surface aeratione
Saccharina latissima SSF Laminarinase, 32 °C, 36 h Saccharomyces cerevisiae (Ethanol 0.45% (v/v) [111]
Red), 32 °C, 36 h
Laminaria digitata2 SSF 2 M HCl + 0.2 M HCl and laminarinase, 24 °C, 69 h Pichia angophorae (CBS 5830), 24 °C, 167 mL/kg algae [50]
69 h
d
Enteromorpha SHF 75 mM H2SO4, 121 °C, 60 min . Celluclast 1.5 L and Saccharomyces cerevisiae (KCTC 8.6 g/L [112]
intestinalis Viscozyme L, 55 °C, 54 h 1126), 30 °C, 48 h
Enteromorpha SSF 75 mM H2SO4, 121 °C, 60 min Celluclast 1.5 L and Saccharomyces cerevisiae (KCTC 7.6 g/L [112]
intestinalis Viscozyme L, 30 °C, 36 h 1126), 30 °C, 36 h
Sargassum spp. SHF 4% H2SO4, 115 °C, 1.5 h. Cellulase and cellobiase, 50 °C, Saccharomyces cerevisiae, 40 °C, 48 h 2.79 g/L [113]
96 h
Gracilaria verrucosa SHF Cellulase and cellobiase, 50 °C, 36 h Saccharomyces cerevisiae (HAU), 0.43 g/g sugars [114]
30 °C, 16 h
3 f
Gelidium amansii SHF 2% H2SO4, 150 °C, 3–3.5 bar, 40 L/h Brettanomyces custersii 27.6/L (0.38 g/ [115]
(KTCT18154P), 30 °C, 39 h g sugar)
Gelidiella acerosa SHF 5% H2SO4 (1:4 ratio), boiling and then room temperature Saccharomyces cerevisiae (MTCC no. 418 mg/g sugars [116]
overnight. 2% cellulase 22086, 45 °C, 36 h 180), 28 °C, 12 h
Gelidium pusillum SHF 5% H2SO4 (1:4 ratio), boiling and then room temperature Saccharomyces cerevisiae (MTCC no. 416 mg/g sugars [116]
overnight. 2% cellulase 22086, 45 °C, 36 h 180), 28 °C, 12 h
Gracilaria dura SHF 5% H2SO4 (1:4 ratio), boiling and then room temperature Saccharomyces cerevisiae (MTCC no. 411 mg/g sugars [116]
overnight. 2% cellulase 22086, 45 °C, 36 h 180), 28 °C, 12 h
4
Saccharina japonica CBP Engineered Escherichia coli (BAL1611), 25–30 °C, 150 hg 4.7% v/v (0.281 g/g [23]
dry algae)

Air rate of 100 mL/min corresponding to an oxygen transfer coefficient (kLa) of 0.053 1/min
SHF: Separate Hydrolysis and Fermentation; SSF: Simultaneous Saccharification and Fermentation; CBP: Consolidated Bioprocessing
M: molar (mol/L); N: equivalent concentration (=M/f_eq with f_eq the equivalence factor)
a
100g seaweed gel coming from 25g of algae.
b
Combination of acid hydrolysis and enzymatic hydrolysis.
c
Successive saccharification with enzyme: two steps.
d
Acid hydrolysis first and then simultaneous enzymatic hydrolysis and fermentation.
e
Oxygen rate is 5.9 mmol O2 /L/h.
f
Continuous hydrolysis reactor.
g
The engineered platform simultaneously allows hydrolysis and fermentation.
1
Collected on a reef.
2
Samples were harvested from wild stock.
3
No origin cited.
4
NO origin cited.

Table 3
Macroalgal conversion into butanol: pretreatment, hydrolysis and fermentation processes. (Reports from 2010 till 2015).

Type of algae Hydrolysis/pretreatment Fermentation Yield (butanol) Reference

Ulva lactuca 1% H2SO4, 125 °C, 30 min Clostridium beijerinckii. Clostridium 0.29 g/g sugar [20]
saccharoperbutylacetonicuma
Saccharina spp. Water pH 2, 65°C, 1 h Clostridium acetobutylicum (ATCC 824), 209 h 0.12 g/g [117]
Ulva lactuca Water, 150 °C, 10 min cellulase (GC220), Clostridium beijerinckii (NCIMB 8052), 140 h 3 g/L (0.35 g ABE/g [24]
50 °C, 24 h sugar)

ABE: Acetone Butanol Ethanol

a
Two-step fermentation: first with Clostridium beijerinckii and second with Clostridium saccharoperbutylacetonicum.

biomass [23,104]. For ethanol and butanol production, before hydro-
lysis, algae are washed, dried and put into smaller pieces (for instance
via milling, grinding, comminution, chopping, beating or extrusion).
Pretreatment and hydrolysis steps aim to improve and fasten the
conversion process. For ethanol and butanol production, in most of the
cases, the biomass undergoes thermal acidic or enzymatic hydrolysis,
which produces sugars from carbohydrates. These sugars are then
transformed into ethanol or butanol through fermentation. The
fermentation organisms are mostly yeasts, but bacteria can also be
used. An interesting completely engineered platform allowing both
hydrolysis and fermentation has been developed using synthetic
biology tools by [23].

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F. Fernand et al.
Table 4
Macroalgal conversion into biodiesel: pretreatment, extraction and transesterification. (Reports from 2010 till 2015).
Type of algae Pretreatment
Ulva lactuca Dried, ground
Asparagopsis taxiformis Dried, ground
Mix of Pelvetia Dried, crushed
canaliculata, Fucus
spiralis
Enteromorpha compressa Dried, pulverized, sieved. 1.5% H2SO4, methanol-oil ratio 12:1, 400 rpm, 60 °C,
Ultrasonication, 24 kHz, 50 °C, 90 min 1% NaOH, methanol-oil ratio 9:1, 600 rpm,
5 min 60 °C, 70 minb
Himanthalia elongata Dried, crushed Algae-methanol ratio (1:15, w/v), NaOH (2 wt%), microwave (800W),
3 mina
FAME: Fatty Acid Methyl Esters
a
Simultaneous extraction and transesterification.
b
Two steps: acid esterification followed by base transesterification.
5. Technologies for algal oil extraction and bioenergy
conversion
In the case of biodiesel production the process is very different.
There is an interesting relation between the harsh extraction proce-
dures and the metabolite profile. The softer the extraction procedure,
the less affected will be the chemical structures [118]. In general the
lipid fraction in macroalgae first undergoes homogenization and
extraction. A couple of extraction protocols have been developed based
on solvents or supercritical fluids as well as on mechanical (i.e. by
ultrasonication) or biological (i.e. by cell wall degrading enzymes)
approaches. Base-catalyzed transesterification transforms fatty acids
derived from triglycerides, phospholipids and galactolipids but also
free fatty acids into the respective methyl esters for further biodiesel
production. As significant amounts of fatty acids might be transformed
into breakdown products upon cell disruption as shown in several
macroalgae, extraction and transesterification can also be achieved
simultaneously in a one-pot reaction [119]. The distribution of fatty
acids in lipids between brown, red and green algae differs strongly, and
variations within the genus level of, for example, Ulva can be observed
and have to be considered as well [120,121]. The biodiesel yield from
macroalgae is thus strongly species dependent.
6. Local perspectives on biofuels production from offshore
cultivated macroalgae
What do data in Tables 1–4 tell us? The production of the biomass
for biofuels is possible, but requires further development of technol-
ogies for both cultivation and processing. Next, based on this global
experience with the offshore biomass to bioenergy production we
analyzed the hypothetical scenario of offshore production of biomass
for transportation biofuels for several countries using currently avail-
able cultivation and processing methods.
In France, according to the National Institute of Statistics and
Economic Studies, in 2013 the total consumption of oil for transporta-
tion was 44.8 million tons of oil equivalents. Since 1 ton of bioethanol
is equivalent to 0.64 ton of oil equivalent (toe), using algae the amount
needed is 70 million tons of bioethanol. Setting a target value of 60%
biofuels with a productivity of 7 g dry weight (DW) of algal biomass per
square meter per day and the possibility to convert 30% of the algal
biomass into transportation biofuel, the marine area needed to cover
(the 60%) of the total oil consumption for transportation in France is
54,795 km2. In comparison, the total French exclusive economic zone
(EEZ) with overseas territories included is about 11 million km2.
For Israel, according to the Central Bureau of Statistics, the total
consumption of transportation fuels in 2013 was 2.8 million toe
Renewable and Sustainable Energy Reviews xx (xxxx) xxxx–xxxx
Extraction/transesterification Oil content Yield of methyl Reference
% DW esters
Chloroform/methanol (2:1, v/v), 6 h. NaOH, methanol, 4.2 3.8% from oil [122]
16 h content
Chloroform/methanol (2:1, v/v), 6 h. NaOH, methanol, 4.1 3.64% from oil [122]
16 h content
Methanol-oil molar ratio 300:1, 1% mass g NaOH/g 2 11.42% from oil [119]
algae, 2.5 L n-hexane/kg algae, 60 °C, 11 ha content
11.14 90.6% from oil [123]
content
2.9 0.034% directly [124]
from dry biomass
(mtoe). Previous studies on the offshore macroalgae cultivation re-
ported a production of 3–25 g DW m-2 d-1 [9,10]. Studies show the
possibility to convert up to 30% of the algal biomass into transportation
biofuels (for example bioethanol) [8–11,13]. To provide the 60% of the
Israel transportation fuels needs, it is required to provide 1665.600
tons of fossil fuels and 2498.400 tons of bioethanol in gasoline gallon
equivalent. Assuming the productivity of 7 g DW m-2 d-1 and a
conversion rate of 0.3, the offshore biomass cultivation area required
to supply 60% of Israel transportation fuels demands is 3250 km2. In
comparison, the total Israel EEZ is 27,000 km2. While Israel has been a
pioneer in land-based macroalgae cultivation [125], there are virtually
no reports of macroalgal offshore culture, except for one in which
Gracilaria and Ulva were tested in nets attached to a fish cage system
ca. 3 miles offshore [126] (Fig. 5a–f). In this short-term series of
experiments both species were cultivated downstream of the fish cages
to study their nutrient uptake, growth and chemical constituents
affected by nutrients derived from the fish. Growth rates could be as
high as 15% per day and consistent levels of fermentable carbohydrates
and proteins were obtained over a range of several weeks [126]. The
generally rough offshore conditions present in the Israeli
Mediterranean shores has evidently preclude further experimental or
pilot efforts to cultivate macroalgae in the sea with currently available
technology. Nevertheless, the high natural irradiance and relatively
warm seawater temperatures year around, in addition to emerging
offshore techniques could encourage further development of cultivation
technology.
The unfavorable environmental conditions along the coasts prevent
to employ conventional techniques [71]. Recently a stakeholder’s
workshop confirmed that there is a clear need, and willingness on
behalf of the policy makers and the research community, to find
sustainable, resource- and space-efficient solutions for combined ocean
use and producing energy is one the possible pathways to use oceans
[127]. Mass culture of benthic macroalgae under rough offshore
conditions in, for example, the North Sea requires rigid culture support
systems that cannot only withstand rough weather conditions but can
also be effectively handled while at the same time retain the cultured
species [26]. In Germany, the pilot studies by Buck and colleagues
demonstrated that S. latissima grew on all of these carriers (Fig. 2C)
with initially up to 14.5% per day and later decreasing length
increments.
Due to the shallow costal sea in Germany, the high sediment load
and harsh conditions, offshore cultivation of brown seaweed S.
latissima was favored and has been successfully performed for fuel
and food production. Moreover various restrictions will not allow an
enhancement of aquaculture production in German coastal regions.
Stakeholders of the coastal regions along with the requirements of the
6
F. Fernand et al.
Fig. 5. Offshore cultivation. (a) Map of the study area showing the fish farm location in Israel. (b) Scheme of the fish cage and the algal culture cages. (c) Location of the algal culture
sites. (d) Scheme of an algal culture cage. (e) Ulva rigida in culture cages attached to the fish cage in Israel. (f) Gracilaria bursa-pastoris in culture cages attached to the fish cage From
[138] with permission. (g). Floater with Saccharina latissima attached to ropes during winter 2014–2015, on the North Sea test-site,10 km West of the island of Texel. This side is
managed by the North Sea Farm foundation. © Royal Netherlands Institute for Sea Research, Nicolas Forjahn, April 2015.
distinguished Marine Protected Areas (MPAs) are limiting the possi-
bilities of aquacultures in German coastal areas. To enhance and
upscale the aquaculture production in German territorial waters a
move away from the coastline to more exposed offshore regions is
necessary. In the North Sea there are several native species which can
be generally considered for offshore cultivations including species from
the Laminariales and Ulvales orders . Preferred areas were identified in
the past for offshore wind farms which are now approved or under
constructions in the North Sea and Baltic Sea (http://www.bsh.de/en/
Marine_uses/Industry/CONTIS_maps/index.jsp). In Germany, the
total transportation energy consumption in 2012 was 56.7 mtons of
toe. Based on the same calculation of the 7 g DW m-2 d-1 of Ulva and a
conversion rate of 0.3 to bioethanol, the total area required to provide
for 60% of German transportation fuels is 69,683 km2. This is almost
twice large than German EEZ which is 33,100 km2.
In the Netherlands the total transportation energy demand in 2012
was 14,2 mtoe. Based on the same calculation of the 7 g DW m-2 d-1 of
Ulva and a conversion rate of 0.3 to bioethanol, the total area required
to provide for 60% of Dutch transportation fuels is 17,368 km2. This is
in comparison to 154,011 km2 of EEZ of the Netherlands. However, in
the Netherlands, offshore cultivation of macroalgae is in its infancy.
Some trials were done before 2014, all with very limited success. In the
winter of 2014–2015, the “kilo-test” was performed, coordinated by the
North Sea Farm foundation. Circular structures were used with a
floater in the middle (Fig. 5g). Ultimately a harvest of 16 kg of fresh
weight S. latissima was achieved. Bad weather conditions, high current
velocities might have caused sub-optimal results. In the Netherlands,
investment cost for offshore seaweed cultivation are estimated to be
€25,000 per ha, while expecting maximum yields of 25–35 tons (DW)
per ha [128]. Total production costs, excluding harvesting and trans-
port are estimated between €1.000- and €1500 per ton DW [129]. It is
clear that it is too early to draw a final conclusion on the economic
viability of developing a North Sea seaweed value chain (as it does not
exist). NIOZ has used the ERSEM model (European Regional Seas
Ecosystem Model, [130]) to incorporate seaweed cultivation in the
North Sea ecosystem. Physiological data for several native seaweed
species was included. The results show that cultivation can take place
with estimated yield of 15–18 ton DW ha-1 year-1. Clearly the location
of cultivation matters. Too far offshore the nutrient concentrations (N
and P) will be lower, resulting in slower growth by the seaweeds.
Similarly the presence of micro-algae matters. The ERSEM model
results show that as soon as the spring bloom of micro-algae starts, the
growth of seaweeds slows down.
In Norway, sugar kelp S. latissima is a species of choice for offshore
cultivation. The yearly outcome from cultivating sugar kelp S. latissima
is predicted to be around 26 tons dry matter ha-1 cultivation area [101].
7
Renewable and Sustainable Energy Reviews xx (xxxx) xxxx–xxxx
Provided 15% dry matter and that 60% of the dry weight is carbohy-
drates this corresponds to 1530 tons of carbohydrates per km2.
Utilization of the three main polysaccharides alginate, mannitol and
laminaran for ethanol production with 80% of the theoretical ethanol
yield [23] would provide 580 tons ethanol per km2. Norway has one of
Europe’s largest economic zones, in total 787,000 km2, and with a sea
area of 125,000 km2 inside the sea boundary. The gasoline consump-
tion in Norway is around 1 mill ton per year, and substituting 60% of
this with bioethanol from sugar kelp would require an area of 1
034 km2, which is less than 1% of the area inside the sea boundary. For
comparison the kelp forest at the Norwegian coast is estimated to cover
10,000 km2. To use the whole biomass or even only the carbohydrate
fraction for biofuel production is not profitable with today's cultivation
technologies, but by extracting valuable fractions first, like alginate and
pigments, the residuals may be an attractive feedstock for biofuel
production due to the possibilities for access to very large amounts.
Progress has recently been made on integrated ethanol and protein
production from kelp biomass, combined with extraction of a high
value compound [131].
7. Environmental and social impacts of offshore macroalgae
cultivation
Addition aspect of offshore macroalgae cultivation is their environ-
mental impact. Macroalgae could impact their surroundings by mod-
ifying biodiversity since there will be competition with other (native)
species [39], hence imposing an environmental threat. Indeed, the risk
of proliferation and invasion of non-native species has been referred in
the past [40–42]. Even if local species are used, massive cultivation of
biomass provides a source for appearance of new mutants, which could
be better fit to the environment than the original population. At the
same time, macroalagae provide a unique environment for living
organisms that supports and encourages biodiversity [39,43,44], which
can also turn into a problem when harvesting this habitat and
damaging the ecosystem [45–51]. In addition, it may worth be
mentioned that in shallow environments a decrease in irradiance
within the water column below seaweed culture settings could have a
negative impact on other marine organisms [52].
However, during photosynthesis and subsequent biomass accumu-
lation, macroalgae will capture significant amounts of inorganic
carbon, first as dissolved CO2 and later as HCO3- after conversion to
CO2. Simultaneously with carbon trapping, photorespiration and
respiration processes will retrieve part of this carbon back into the
seawater. A portion recombines into CO2 through biodegradation while
the rest stays as Particulated Organic Carbon (POC) in the seawater
column, and finally deposits in the sea bottom. Since POC turnover is
F. Fernand et al.
in the order of 100–1000 years, this process would serve as a carbon
sequestration, at least for some time [39].
In addition to inorganic carbon, macroalgae also use other inor-
ganic nutrients to grow, such as nitrogen and phosphorous. In
eutrophic waters with excessive nutrients, a wild over-development
(bloom) of macroalgae may take place. The generally nuisance blooms
can be visible in coastal waters [85–90] and disturbs the ecosystem in
place [42] and can eventually lead to hypoxia and to the reduction of
aquatic biodiversity [91]. It has been noted that the impact is reduced
in deep waters [17]. Thus, controlled macroalgae cultivation can be
used for bioremediation [61,86,90], i.e. scavenging of pollutants
(inorganic nutrients in this case) dissolved in seawater, or heavy metals
[92–94]. The excess of nutrients can be removed while harvesting of
the macroalgae [86,95]. For example, macroalgae cultures have been
deployed near fish farming cages (Fig. 5a–f) [96,97] or for corals
protection [98].
Nutrients derived from fish cultivation can be used by seaweeds in
integrated multi-trophic aquaculture (IMTA), as fertilizers for algae in
both land based or offshore marine culture systems [99,100]. Timing
and tuning of the flexible fluxes during the year are essential and hard
to achieve. For example, the main growth season for a kelp is in the
spring whereas the effluents from salmon farms are highest in the late
summer and early autumn [83]. By modeling it has been shown that a
kelp farm of 1 hectare will take up less than 0.5% of the total amount of
dissolved inorganic nitrogen from a 5000 tons salmon farm, indicating
that large cultivation areas would be needed to consume all the
dissolved nutrients that are released from the fish farms [101].
Macroalgae cultivated at sea can be regarded as last site where the
scarce resource phosphate can be captured. In spite of intensive efforts
on land and in freshwater bodies, still considerable amounts of
phosphate are washed to sea and often multiple regulatory issues of
the stakeholders have to be sorted out. Macroalgae offer the last
opportunity to capture this phosphate before it is further and further
diluted in (deep) ocean waters. Therefore, mass cultivation of commer-
cially valuable seaweeds plays an increasingly important role as
nutrient removal-system [102].
Concerning the possible social impacts, marine biorefineries could
transform the economies of the low income farmers, as the major
seaweed cultivation today is done by on the family level poorest
farmers, mostly in Indonesia, Philippines, China, India and Tanzania,
who earn US$0.5 - 3 a day [103]. The macroalgae are cultivated
offshore and are sold fresh, dried and semidried for processing, mostly
for food ingredients industry. The cost of macroalgae at the farm today
varies from $70 to $150 dry weight ton, while the final extracted
products are sold for an order of magnitude more [103]. Moreover,
only 8–12% of the cultivated biomass is used for the food chemical
production, the rest is disposed as waste. Development of new
technologies for cultivation in the near-future explorable areas, and
macroalgae processing to platform chemicals have a potential to
provide additional income to the thousands of families who are already
involved in macroalgae farming [103].
8. Challenges ahead
Macroalgae are very promising, yet very challenging source of
biomass for bioenergy biorefineries when it comes to massive cultiva-
tion. Major future effort should be invested in the fundamental biology
of macroalgae to allow for effective breeding and reproduction meth-
ods. High-throughput strains selection methods, possibly based on
imaging as shown for plants phenomics [132] could transform the
traditional approaches for breeding and selection. At the same time,
information on the interaction of the selected species with environ-
ment, their invasiveness and incorporation into the local and global
food webs will be required before release to the offshore environment.
In most of the regions where kelp is cultivated the biomass is
normally harvested in the late spring or early summer, due to high
8
Renewable and Sustainable Energy Reviews xx (xxxx) xxxx–xxxx
water temperatures or biofouling problems during the summer. The
landing of several hundred tons of wet kelp over a restricted number of
weeks claims for good logistic solutions regarding transportation,
stabilization and storage of the biomass, or development of industrial
scaled processing lines ready for continuous extraction and conversion
of the biomass as it enters the factory, whether this is on a vessel or on
land.
Cultivation offshore opens new opportunities to produce sustain-
able biomass without using more arable land and freshwater. Yet,
harsh offshore environment currently is challenging to work in on the
large areas and novel technologies for automatic cultivation and
harvesting are needed. Automation transformed the terrestrial agri-
cultural sector allowing for more efficient resource use. Macroalgae
cultivation today is mostly traditional and requires a lot of unqualified,
low paid hand labor [103]. Intensified and automated cultivation could
provide a new job sector that could contribute to sustainable develop-
ment in many rural areas. Although the areas for the biomass
production are large, future development of cultivation technologies,
for example with intensification of offshore growth [133], or more
complete conversion of macroalgae biomass, could significantly reduce
the areas and required process costs. Massive offshore cultivation of
macroalgae will consume large amounts of nutrients. As based on
naturally occurring nutrient levels only, year round cultivation which is
often required for a commercial facility, will likely be non-sustainable.
Pumping up of nutrient-rich, deep seawater layers (artificial upwelling
systems) could provide a solution for sustainable fertilization [134–
136]. However, the technologies for artificial upwelling are still in the
early stages of development and their impact on the natural environ-
ment is still poorly understood.
On the conversion aspects, macroalgae chemical composition is
complex. Different from terrestrial biomass, where the primary carbo-
hydrate is glucose, macroalgal biomass has multiple “unusual” carbo-
hydrates such as galactose, rhamnose and arabinose and sugar acids,
which are difficult to ferment with conventional microorganisms. Novel
microorganism, from marine environments, should be identified and
domesticated for complete conversion of marine biomass into useful
bioenergy molecules. Importantly, the information about net energy
return of investment and economics of offshore biomass for bioenergy
production and conversion is not known. Although projections can be
made based on the pilot studies and experience gains from offshore oil,
wind and gas production, cost of installation and particularly main-
tenance of large areas required for biomass production are challenging
for economic estimations. Data that combine energetic, economic and
environmental efficiency of offshore bioenergy systems can be analyzed
using emerging methods of energy systems modeling such as environ-
mental exergenomics [137]. Environmental exergonomics allows for
analysis and optimization of energy systems using technological,
economic, and environmental exergy flows by maximizing the system
energetic and economic outputs but minimizing the damages to eco-
systems.
Acknowledgements
FF, AI and AG thank TAU Center for Innovation in Transportation
and the Ministry of National Infrastructure, energy and Water
Resources for the support of this study, AI thanks the Ministry of
Science, Technology and Space, Israel, for their support through Grant
no. 3-99763. All authors would like to acknowledge networking
support by the COST Action “Phycomorph” FA1406.
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