Author Version of : Carbohydrate Polymers, vol.

267; 2021; Article no: 118241

Seaweed-based cellulose: applications, and future perspectives

Ravi S. Baghel1*, CRK Reddy2 & Ravindra Pal Singh3

1
Biological Oceanographic Division, CSIR-National Institute of Oceanography,
Dona Paula, Goa, India 403004
2
Indian Centre for Climate and Societal Impact Research
Vivekanand Research and Training Institute, Mandvi-Katch, Gujarat 370465
3
Food and Nutritional Biotechnology Division, National Agri-Food Biotechnology Institute (NABI),
SAS Nagar, Punjab, 140306, India.
Corresponding author Email: [email protected]; [email protected]
Phone: +91 8322450295 Fax: +91 8322450606

Abstract
Cellulose is a naturally occurring organic polymer extracted mainly from lignocellulosic biomass of
terrestrial origin. However, the increasing production of seaweeds for growing global market demands
has developed the opportunity to use it as an additional cellulose source. This review aims to prepare
comprehensive information to understand seaweed cellulose and its possible applications better. This is
the first review that summarizes and discusses the cellulose from all three types (green, red, and brown)
of seaweeds in various aspects such as contents, extraction strategies, and cellulose-based products. The
seaweed cellulose applications and future perspectives are also discussed. Several seaweed species were
found to have significant cellulose content (9-34% dry weight). The review highlights that the properties
of seaweed cellulose-based products were comparable to products prepared from plant-based cellulose.
Overall, this work demonstrates that cellulose could be economically extracted from phycocolloids
industrial waste and selected cellulose-rich seaweed species for various commercial applications.
Keywords: Bioethanol, Cellulose, Seaweed, Microcrystalline cellulose, Nanocellolose

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1. Introduction
Cellulose is the most common biomaterial derived from renewable resources such as terrestrial plants,
algae, and fungi (Gupta et al., 2019). Structurally, cellulose is a homogeneous polymer made up of β- (1,
4) linked β-D-glucopyranose units (Moon et al., 2011). Cellulose has been attracting researchers' attention
for a long time and is used extensively in various applications due to its abundant availability, low cost,
and easier processability (Motaung & Linganiso, 2018). Cellulose versatile characteristics such as low
weight, non-toxicity, mechanical strength, hydrophilic and hygroscopic nature, biodegradability, and
recyclability make it suitable for various commercial applications (Hafid et al., 2021; Zhang et al., 2021;
Du et al., 2018). Cellulose and its derivatives such as cellulose ester, cellulose ethers, cellulose fiber,
microcrystalline cellulose, and nanocellulose are widely used in paper, textiles, pharmaceuticals,
veterinary, cosmetic, food industries, and water treatment (Kassab et al., 2020; He et al., 2020; Arca et
al., 2018; Lakshmi et al., 2017). The annual production of cellulose is estimated at 180 billion tons
(Sundarraj & Ranganathan, 2018) and, the market demand for cellulose and its derivates is steadily
increasing (Hafid et al., 2021; Zhang et al., 2021). The cellulose industries highly depend on
lignocellulosic biomass like wood, cotton, flax, hemp, and Jute for cellulose production (Lavanya et al.,
2011). The removal of lignin is the major challenge in extracting cellulose from lignocellulosic biomass
(Lee et al., 2014). The harsh chemical treatment to biomass leads to partial degradation of the cellulose
(Ververis et al., 2004) and generates effluents with severe environmental concerns. Therefore, easily
available and fast-growing biomass, such as seaweeds, are also gaining global attention as an additional
source of cellulose.
Seaweeds (Macroalge) are broadly classified into Chlorophyta, Rhodophyta, and Phaeophyta, based
on their pigmentation (Samiee et al., 2019; Kraan, 2013). Chlorophyta consists of green seaweeds that
contain chlorophyll a and b as a primary pigment. The Rhodophyta is a group of red seaweeds that contains
phycobiliprotein dominantly. The third is Phaeophyta (brown seaweed) contains fucoxanthin dominantly.
Seaweeds, being aquatic organisms, contain copious amounts of water in their bodies up to 85% (Badmus
et al., 2019), while the rest is represented mainly by organic matter and minerals. The dry matter of
seaweeds comprises total lipids 0.5-3.5% (Kumari et al., 2010), proteins 3-50% (Harnedy and FitzGerald,
2011), total carbohydrates 21-61%, and minerals 12-46% (Baghel et al., 2014; Kumar et al., 2011). A
higher growth rate, fewer inputs in seaweed cultivation (require no land and fertilizers, pesticides, and
freshwater) makes them an attractive option over the terrestrial biomass (Baghel et al., 2017). Owing to
this reason, the world production of seaweeds increased by three times in 2018 (32.4 million tons) from
10.6 million tonnes in 2000 (FAO, 2020). Of total seaweed harvest, 97.1% volume came from farmed
seaweeds and 2.9% from wild harvest. Certain seaweeds have been utilized in many countries like Costa
Rica, Japan, China, and Egypt from prehistoric times as a human diet (Dillehay et al., 2008). The red
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seaweed Porphyra (Nori) is used to prepare sushi in various countries like Japan, South Korea, United
States, and the United Kingdom due to its richness in proteins and vitamins (Merrill, 1993; Chapman,
2013). The phycocolloids (Carbohydrates) such as agar, carrageenan, and alginate are the main
commercial products traditionally extracted from seaweed biomass (Khalil et al., 2018). The hydrocolloid
yielding seaweeds are divided into three groups. The agar-bearing seaweeds are known as agarophytes,
carrageenan-bearing seaweeds are known as carrageenophytes, while alginate-bearing seaweeds are called
alginophytes (McHugh 1991). These hydrocolloids are extensively used as an ingredient in various
industries ranging from food, beverages, dairy, animal feed, cosmetics, personal care, pharma, healthcare,
textile, printing, and paper coating (Baghel et al., 2017). Also, the genus Ulva and Porphyra members
contain significant amounts of storage sulfated polysaccharides such as ulvan and porphyran. Such
polysaccharides exhibit various biological activities with the potentials to develop drugs and
nutraceuticals (Pereira, 2020). The seaweed-based minerals are yet other components commercially used
as fertilizer (Baghel et al., 2020). Apart from these, seaweeds have very little or no lignin helping to
recover pure and intact cellulose with greater suitability for biomedical applications (Halib et al., 2017).

The research on seaweed cellulose has begun long ago. Cellulose from green seaweed was first
described in 1885 (Mihranyan, 2011). Cellulose is present in the seaweed cell wall with other diverse
polysaccharides such as xylan, mannan, galactan, alginic acid, agar, carrageenan, rhamnose-
glucuronate/iduronic; depending on the type of the seaweed groups (Wahlström et al., 2020a; Ciancia et
al., 2020; Michel et al., 2010). Chlorophyceae members are known to have a wide array of walls ranging
from cellulose–pectin complexes to ones made of hydroxyproline-rich glycoproteins (Domozych et al.,
2012). Figure 1 describing the cell walls of green, red, and brown seaweeds where cellulose is present in
the cell wall as the fibrillar skeleton material. In green seaweed, such as Ulva species- the cell wall is
composed of four polysaccharides families. Amid them, water-soluble ulvan and insoluble cellulose as
major constituents, and certain hemicellulose constituents such as β-1,4-D-glucuronan and β-1,4-D-
xyloglucan present in minor fractions (Fig. 1A) (Lahaye and Robic, 2007). Further, the most abundant
fibrillar constituents of the cell walls of Ulvophycean may change from cellulose to β-mannans to β-xylans
during different life cycle phases (Domozych et al., 2012). Red seaweed cell walls made up of cellulose
microfibrils, glucomannan, and hemicellulose material, consisted of sulfated glucan and xylogalactans
(Fig. 1B) (Lechat, 1998). For instance, β-D-(1→4), β-D-(1→3) mixed linkage xylans are known from
Nemaliales and Palmariales (Viana et al., 2011). Sulfated galactans are another major sugars present in
the cell walls of red seaweeds, present in the form of carrageenan and agar (Usov, 1998). These xylans
and galactans are predicted to be interconnected in cell walls (Fig. 1B); however, proper structures are yet
to be elusive. Contrary, the fucose-containing sulfated polysaccharides (such as fucoidans) are the major
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crosslinking glycan present in cell walls of brown seaweed (Fig. 1C). These sulfated polysaccharides play
a role in interlocking the cellulose microfibrils in layers parallel to the cell surface that create the
framework of the cells (Deniaud-Bouët et al., 2017; Michel et al., 2010). Another peculiar feature of brown
seaweed is alginate, which can be present either in the homopolymer form of d-mannuronic acid /l-
guluronic acid or present in an alternative fashion (Fig. 1C). Notably, lignin is either absent in seaweeds
(Ge et al., 2011) or present in very low concentrations (Wi et al., 2009) compared to land plants.

Figure 1 Cell wall composition of seaweeds; 1A, representative cell wall for green seaweeds; 1B,
representative cell wall for red seaweeds and 1C, cell wall composition of brown seaweeds. (adapted
and modified from Stiger-Pouvreau et al., 2016: Michel et al., 2010; Lahaye and Robic, 2007; Lechat,
1998)

The cellulose sourced from different seaweeds species reported to have similarities with cotton
cellulose based on three different analysis, i.e., 1) production of D-glucose only upon hydrolysis, 2)
presence of β- (1, 4) linkage confirmed by the preparation of cellobiose octa-acetate and 3) similar X-Ray
diagram from algal cellulose and cotton cellulose (Percival& Ross, 1949). However, algal cellulose has
differed from cotton/plant cellulose in the compositional ratio of two allomorphs. The algal cellulose is
dominated by α allomorph (α cellulose) (Siddhanta et al., 2009) while cotton type cellulose dominated by

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β allomorph (β cellulose) (Tsekos, 2002; Horii et al., 1987). Especially the cellulose content and allomorph
were analysed in 12 seaweed species belonging to green, red, and brown seaweeds (Fig. 2) (Siddhanta et
al., 2009). They found that α allomorph was predominant in all studied species. Interestingly, it showed
that species belongs to the agarophytic and alginophytic group contained high cellulose and α-cellulose
contents, while the member of carrageenophyte having low cellulose. The dominance of α allomorph in
seaweed cellulose makes them more thermodynamically reactive than woody or plant biomass-based
cellulose (Siddhanta et al., 2009). Apart from that, seaweed cellulose and plant cellulose reported having
a different surface area. Plant cellulose has a specific surface area of 1 square meter per gram; on the
contrary, seaweed, Cladophora having up to 100 times larger surface area as algal cellulose extruded as
single strands (Marsin et al., 2005).

Green seaweeds classified into three groups based on their cell wall constituents (Nicolai &
Preston,1952). Groups 1 comprised the green algae belonging to the two orders, Cladophorales
(Cladophora, Chaetomorpha, Rhizoclonium, and Microdyction) and Siphonocladales (Valonia,
Dictyosphaeria, Siphonocladus, and Boergesenia), having the high crystalline native cellulose as the
major component of cell walls. The bulk of green algae with a large quantity of mercerized like low
crystalline randomly oriented cellulose in their cell walls are classified as Group 2. The small group of
heterogeneous algae, in which case cellulose is a minor component in their cell walls like Vaucheria and
Spirogyra included in Group 3. The crystalline features were analyzed for algal celluloses extracted from
different species (Koyama et al., 1997), which reported three types of cellulose in the algal system: 1) Iα-
rich, broad microfibril, 0.6 nm oriented type; 2) Iβ-dominant, flat-ribbon, 0.53 nm oriented type; and 3)
Iβ-dominant, small, random oriented type. The agarophytic seaweeds were reported to have higher
cellulose content than carrageenophytes (Siddhanta et al., 2009). Further, Siddhanta et al. (2011) studied
cellulose content and its allomorph for 21 seaweed species belonging to the different classes from Indian
waters. They established a relationship between the morphology of algal thallus and cellulose yields. The
algal species with compact morphology have higher cellulose content than succulent morphology.

The properties of seaweed-based cellulose vary with the source species. The cellulose fiber extracted
from various seaweed species reported having different lengths and widths. For example, fiber from
Gelidium (red seaweed) was reported with 0.5-1.0 mm length and 3-7 μm width based on scanning
electron microscopy images (Seo et al., 2010a, b). Green seaweed Cladophora cellulose was reported
with 0.7-2.3 mm length and 1.3-1.6 μm width (Leopold & Marton, 1974). The cellulose fiber extracted
from another green seaweed Rhizoclonium was having 0.2-3.3 mm fiber length (Chao et al., 2000).
Similarly, there has also been a difference in the crystallinity index of the cellulose from seaweeds. For
example, Cladophora cellulose had 84% (Hallac & Ragauskas, 2011); 67.0% for Kelp cellulose (Liu et

5
al., 2017a), and 48 % for Ulva lactuca (Wahlström et al., 2020b). Further, Cladophora based cellulose
was reported to have a high degree of polymerization (DP) with a weighted average DP of about 2,000
(Hallac& Ragauskas, 2011). The subsequent studies have also investigated various seaweeds to quantify
the cellulose yield (Chen et al., 2016; Baghel et al., 2014; Siddhanta et al., 2013, 2011, 2009; Mihranyan,
2011) (Table 1,2,3).

Figure 2 Yields (%) of cellulose, α- and β-cellulose for different seaweed species. (adapted from
Siddhanta et al., 2009)

Due to the high ever-growing demands of cellulose and its derivatives, it is essential to identify the
additional sources of cellulose along with a sustainable and efficient extraction process (Hafid et al., 2021).
In the past decades, seaweeds have been mainly processed to extract hydrocolloids as the major known
constituents of biomass (Baghel et al., 2014). However, the last decade has witnessed significant
advancements in developing innovative biomass processing technologies facilitating the successful
extraction of cellulose from residual biomass leftover following the extraction of minerals, protein, and
hydrocolloid (Baghel et al., 2015, Trivedi et al., 2016). The preparation of cellulose from such residual
biomass has provided an opportunity for the large-scale extraction of cellulose and its utilization for
various applications. With this backdrop, the review is begun with the general introduction of seaweeds
and their uses. The presence of cellulose in seaweed cell walls and its similarities and differences with
plant-based cellulose are described. Further, cellulose contents in various seaweed species are

6
summarized. The cellulose extraction strategies are discussed with examples and compared with the
cellulose extraction process from lignocellulosic biomass. The standing stock of cellulose in annual
phycocolloids industries waste is estimated. Further, seaweed cellulose-based products and their
application are reviewed and also compared with products prepared from plant-based cellulose. At last
concluding remarks and an outline for future research, programs are provided.

2. Cellulose contents in seaweed

Several species belonging to the green, red, and brown seaweeds have been investigated for their
cellulosic content. A total number of 19 species of green seaweeds were studied for cellulose content that
ranged from 1.5-34% on dry weight (DW) of biomass with an average being 9.67% (Table 1). Whereas
the same in 47 species of red seaweeds ranged from 0.85-18% DW with an average being 4.75% (Table
2). The corresponding value for the brown seaweeds ranged from 2.2-10.2% on a DW basis, with an
average being 7.88% based on the investigation of a total of 15 species (Table 3). Overall the cellulose
content reported for seaweed biomass ranges between 0.85 to 34%. The investigation on cellulose content
from different seaweeds showed that cellulose content not only differs with a different group of seaweed
but also among the members of the same group. The member of red seaweed showed higher diversity in
cellulose content in comparison to green and brown seaweeds. Among the seaweeds investigated, about
27 species contained cellulose in the range of 9-34 % on a DW basis, indicating a potential opportunity
for their commercial cultivation for cellulose production.

Table 1 Cellulose content in green seaweeds on dry weight (DW) basis

S. Order Species Cellulose References

N. (% )
1 Ulvales Ulva fasciata 11-14.7 Trivedi et al., 2016; Bhutiya
et al., 2018., Lakshmi et al.,
2017
2 Ulva lactuca 5.6-16.3 Siddhanta et al., 2011; Doh
et al., 2020a
3 Ulva rigida 7.5 Siddhanta et al., 2011
4 Ulva prolifera 15.3 Gao et al., 2015
5 Ulva compressa 3.5
6 Ulva prolifera 5 Siddhanta et al., 2011
7 Cladophorales Valoniopsis pachynema 6.8
8 Chamaedoris auriculata 9 Siddhanta et al., 2009

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 9 Chaetomorpha antennina 8-34 Siddhanta et al., 2011;
Bhutiya et al., 2018
10 Chaetomorpha aerea 20 Siddhanta et al., 2011
11 Cladophora glomerata 21.6 Xiang et al., 2016
12 Boodlea composite 5.2
13 Bryopsidales Codium dwarkense 1.5
14 Codium tomentosum 7
15 Caulerpa taxifolia 11
16 Caulerpa imbricate 7.4 Siddhanta et al., 2011
17 Bryopsis hypnoides 4.3
18 Halimeda macroloba 3.7
19 Ulotrichales Acrosiphonia orientalis 10.4
Average cellulose content 9.67

Table 2 Cellulose content in red seaweeds on dry weight (DW) basis

S. Order Species Cellulose References
N. (%)
1 Nemaliales Scinaia carnosa 1.2 Siddhanta et al., 2011
2 Liagora ceranoides 0.85
3 Liagora indica 0.9 Siddhanta et al., 2013
4 Galaxaura oblongata 1.8
5 Halymeniales Grateloupia indica 4.2 Siddhanta et al., 2011
6 Grateloupia lithophila 2.9 Siddhanta et al., 2013
7 Grateloupia filicina 2.9
8 Halymenia venusta 3.6
9 Corallinales Cheilosporum spectabile 2.5
10 Gigartinales Solieria robusta 2.1
11 Hypnea valentiae 2
12 Hypnea pannosa 2 Siddhanta et al., 2011
13 Kappaphycus alvarezii 2
14 Sarconema scinaioides 2.1
15 Sarconema filiforme 4

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16 Rhodymeniales Champia parvula 2.1
17 Champia indica 2.25
18 Champia globulifera 2.5 Siddhanta et al., 2013
19 Ceramiales Laurencia majuscule 3.4
20 Ceramium cruciatum 2.4
21 Dasya naccarioides 2.6
22 Neurymenia fraxinifolia 3
23 Nitophyllum marginale 1.4
24 Acanthophora dendroides 3.3
25 Haloplegma duperreyi 3.4 Siddhanta et al., 2011
26 Laurencia nana 3
27 Gracilariales Gracilaria millardetii 3.8
28 Gracilaria pudumadamensis 2 Siddhanta et al., 2013
29 Gracilaria dura 2.4-3.7 Baghel et al., 2014;
Siddhanta et al., 2011
30 Gracilaria edulis 4.88-5.3
31 Gracilaria debilis 4.2
32 Gracilaria textorii 3.6-4.62
33 Gracilaria fergusonii 4.2-4.3
34 Gracilaria salicornia 4.9 Baghel et al., 2014
35 Gracilaria corticata 5.27
36 Gracilaria. c. v. cylindrica 6.08
37 Gracilaria foliifera 3.77
38 Gracilaria crassa 7.21
39 Gracilaria verrucosa 14-18 Kumar et al., 2013
40 Gelidiales Gelidium micropterum 10.63 Baghel et al., 2014
41 Gelidium variabilis 11.38
42 Gelidium amansii 9.09 Gao et al., 2015
43 Gelidium elegans 17.2 Chen et al., 2016
44 Gelidium pusillum 9.3-12.20 Siddhanta et al., 2011;
Baghel et al., 2014
45 Gelidiella acerosa 9.77-13.6 Baghel et al., 2014;
Siddhanta et al., 2011

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 46 Pterocladia heteroplatos 8.7 Siddhanta et al., 2013
47 Bangiales Porphyra umbilicalis 9.84 Gao et al., 2015
Average cellulose content 4.75

Table 3 Cellulose content in brown seaweeds on dry weight (DW) basis

S. Order Species Cellulose References
No. (%)
1 Dictyotales Dictyota dichotoma 9.5 Siddhanta et al., 2011
2 Dictyota bartayresianac 9.3
3 Padina tetrastromatica 9.5
4 Dictyota divaricata 5.2 Siddhanta et al., 2013
5 Padina pavonica 6.1
6 Spatoglossum asperum 2.2
7 Stoechospermum marginatum 4.5
8 Pocockiella variegata 9.5
9 Fucales Hormophysa triquerta 8.2
10 Sargassum tenerrimum 10 Siddhanta et al., 2011
11 Cystoseira indica 9
12 Iyengaria stellata 9.2
13 Laminariales Macrocystis pyrifera 5.9 Gao et al., 2015
14 Saccharina japonica 9.9 He et al., 2018
15 Scytosiphonales Hydroclathrus clathratus 10.2 Siddhanta et al., 2013
Average content 7.88

3. Cellulose extraction: Primary biomass vs residual biomass

Primarily cellulose is extracted from whole seaweed dry biomass (Mihranyan et al.,2004; Siddhanta
et al., 2013, 2011, 2009; Baghel et al., 2014). The cellulose extraction method presented in Figure 3A
includes (1) bleaching of dried seaweed biomass with sodium chlorite (NaClO2) in an acetate buffer at 60
°C for 3-6 hrs followed by washing to make material neutral. (2) Alkali treatment (0.5 M NaOH) at 60
°C for 6-12 hrs and subsequent washing of material till neutral pH. (3) Heating of alkali-treated material
till boiling in 5% HCl (V/V) followed by incubation at room temperature. (4) Washing of material till
neutral pH and drying to get cellulose. However, in recent times, integrated approaches have been made
to extract cellulose from seaweed biomass along with several co-products (Baghel et al., 2020; Trivedi et
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al., 2016; Kumar et al., 2013). For example, 27-35% of cellulose was extracted from residual pulp that
leftover after sequential recovery of various components such as pigments, minerals, lipids, and agar from
three different agarophytic seaweeds Gelidiella acerosa, Gracilaria dura, and Gelidium pusillum (Baghel
et al., 2015). The total cellulose quantity extracted from the respective samples' residual pulp was
comparable with those obtained from primary biomass using conventional methods. The residual pulp of
Gracilaria verrucosa biomass leftover after agar extraction was reported to have about 40% cellulose
(Kumar et al., 2013). In the integrated process, as much as 30% cellulose on a dry weight basis has been
recovered from the residual biomass of the green seaweed Ulva fasciata (Trivedi et al., 2016). More
recently, about 45% of cellulose has been recovered from the residual biomass of sargassum tenerrimum
(Fig. 3B) with minimum chemical inputs and treatments (Baghel et al., 2020). Interestingly, extraction of
seaweed components such as pigments, minerals, proteins, and hydrocolloids before the cellulose
extraction has not affected the cellulose yields; it instead got densified in residual mass with the recovery
of various components in successive steps (Baghel et al., 2020, 2015; Trivedi et al., 2016). The cellulose
remained intact in the residual biomass due to its insoluble nature in the water despite pre-treating the
biomass in several aqueous solutions for recovering other products. The cellulose extracted from residual
seaweed biomass reported having similar characteristic peaks in FTIR analysis than standard Whatman
cellulose. Further, 86-89% saccharification efficiency upon enzymatic hydrolysis confirmed the purity
and quality of cellulose (Baghel et al., 2015; Trivedi et al., 2016).

Cellulose extraction from such cellulose-dense seaweed residual mass helps minimize the chemical
usage and energy expenses in the extraction process compared with cellulose extraction from whole dried
primary seaweed biomass. The cellulose contents (0.85-34% DW) of seaweed are not comparable with
lignocellulosic biomass, ranging from 25-45% (Lee et al., 2014). However, the cellulose recovered from
residual seaweed biomass is well comparable with that of higher plants as it ranges from 27-40% on a
DW basis. The cellulose extraction process for the seaweed's whole biomass and residual biomass was
compared with the process of cellulose extraction from lignocellulosic biomass paddy straw to describe
the number of treatments and inputs needed for extraction (Rezanezhadet al., 2013)(Fig. 3). Paddy is one
of the major crops for Asian countries. Further, paddy straw is the primary low-value agricultural waste.
Especially in India, it is used to burn in the field after crop harvesting, creating severe air pollution; thus,
it is considered a potential alternative cellulose source. However, cellulose extraction from paddy straw
is needed high chemical inputs and harsh treatments (Fig. 3C) such as 3% KOH, 10% HCL, NaClO2
bleaching, 5% NaHSO4 treatment followed by 17.5% NaOH to remove lignin and other components
present in it while compared with seaweed biomass.

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Figure 3 Comparison of cellulose extraction process for seaweed dry biomass; (3A), seaweed residual
mass (3B) and rice straw a lignocellulosic biomass (3C).

4. Cellulose from seaweed phycocolloids industrial waste

The global phycocolloids industries consume 5,94,220 dry tons of seaweed biomass annually to
produce carrageenan, agar, and alginate (Porse & Rudolph 2017), and residual biomass was treated as
waste or without considering for any commercial use (Baghel et al., 2015). These industries mainly using
algal species belonging to the group of red and brown seaweeds. Firstly an average cellulose content of
6.31% was calculated using the data published for cellulose contents in several red and brown seaweeds
(Table 2 & 3) to estimate cellulose quantity present in annually generated seaweed residual biomass from
hydrocolloid industries. Further, based on average cellulose content data, it was estimated that about
37,495 tons of cellulose could be produced annually from residue leftover from hydrocolloid industries
using the formula presented as equation 1. The processing of such industrial waste provides cellulose for
various applications and helps to convert waste to wealth.

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 (1)

Wherein ECQ is Estimated cellulose quantity in hydrocolloids industrial waste (tons); AUB is Annually
utilized biomass in tons, and ACC is Average cellulose content reported for red and brown seaweeds
(Table 2 & 3).

5. Seaweed-cellulose based products

Although seaweed cellulose research has begun long ago, there are still limited research reports
available on its application. Seaweed-based cellulose was widely studied for bioethanol production apart
from some scanty reports for its uses in the preparation of the paper, nanocelluloses, microcrystalline
cellulose, carboxymethyl cellulose, and bioplastic, etc. The seaweed cellulose-based products are
described below in detail.

5.1. Nanocellulose
Nano-celluloses are unique and promising natural materials among the products derived from
cellulose. The nanocelluloses are categorized into three types based on structural properties and source;
(1) nanocrystalline cellulose (CNC), (2) cellulose nanofiber (CNF) and, (3) bacterial cellulose (BC), also
called microbial cellulose (Jawaid et al., 2017). The size and crystallinity are the major characteristics of
nanocelluloses. Nanocelluloses are prepared by hydrolysis of amorphous cellulose regions using sulphuric
acid (Habibi et al., 2010). The nanocellulose has a typical diameter of 10-50 nm and length in nm to
several micrometers (Abe et al., 2007; Jonoobi et al., 2012). The physical and chemical properties of
nanocellulose and unique behavior such as liquid crystals, membranes, gel, and emulsions at the nanoscale
developed an opportunity for its potential application as a soft material in food, cosmetics, and biomedical
applications (Kim et al., 2019; Nystrom et al., 2018; Fatona et al., 2018; Lin & Dufresne, 2014).
Nanocellulose is also used in other sectors such as paper and pulp, sensors, and electronics usages. The
present estimated global nanocellulose market is USD 297 million and is projected to reach USD 783
million by 2025, with a 21.3% compound annual growth rate (www.marketsandmarkets.com).

There has been extensive research done on the preparation of nanocellulose from different
lignocellulosic biomass of terrestrial origin. Mainly pulps such as hardwood pulp, softwood kraft pulp,
recycled pulp, and different fibers like cotton fiber, flax fiber, grass fiber, ramie fiber, sisal fiber are used
to prepare it (Bettaieb et al., 2015). Last decade few reports were published for nanocellulose preparation
either using seaweed biomass or residual seaweed waste. The cellulose nanofibrils were prepared from
cellulose extracted from green seaweed Cladophora glomerata using a two-step treatment. In the first

13
step, cellulose was enzymatically hydrolyzed and in 2nd step subjected to mechanical nanofibrillation
using a domestic blender at 20000 rpm (Xiang et al., 2016). The prepared nanofibrils' size was 10-40 nm
diameter and having a 96.5% crystalline index. Moreover, cellulose nanofibrils showed higher thermal
stability in compassion to CNFs produced from bleached eucalyptus pulp (Xiang et al., 2016). The higher
thermal stability of CNFs may be attributed to their high crystallinity, low bound water, and high purity.
Later various studies have focused on investigating the seaweed-based cellulose for its suitability for
nanocellulose preparation (Table 4). There were two reports wherein the researchers successfully prepared
cellulose nanocrystals from the residual waste of red and brown seaweed biomass (Achaby et al., 2018;
Liu et al., 2017a). Liu et al. (2017a) used kelp waste to prepare NCC with a 52.3 % yield and 69.4%
crystallinity index. Achaby et al. (2018) prepared nanocrystal from red seaweed residues leftover after
agar extraction by H2SO4 hydrolysis for different periods such as 30, 40, and 80 min. The NCC yields and
crystallinity index was recorded in the range of 11.18 to 13.76 % and 81 to 87%, respectively. The 80 min
acidic hydrolysis yielded 11.18% nanocellulose with 87% crystallinity and a diameter of 5.2±2.9 nm (Fig.
4A). Nanocellulose prepared from seaweed waste was superior in yield, crystallinity, and size to that
extracted from whole Gelidium elegans biomass (Chen et al.; 2016) (Fig. 4B).

Figure 4 Comparison of nanocellulose isolated from Gelidium residual waste left after agar extraction
(4A) and whole Gelidium biomass (4B). (Fig. 4A adapted and modified from Achaby et al., 2018; Fig. 4B
adapted and modified from Chen et al., 2016)
14
 Recently, Doh et al. (2020a,b) studied cellulose from many seaweed species for nanocrystal
preparation. The mechanical and chemical treatments were used to prepare nanocrystals having 58-99%
crystallinity index and a length ranging 21-248 nm and width 4.8-41 nm analyzed using a high-resolution
transmission electron microscope (TEM) and measured with Image J software. Overall, nanocellulose
reported from different macroalgal biomass showed a good crystallinity index ranging from 58-99% and
size in diameter ranging from 2.3-40 nm (Table 4). There were three different treatments, chemical,
enzymatic, and mechanical, alone or in combination, used to prepare nanocellulose from seaweed
biomass. The properties of nanocellulose were recorded highly variable with source seaweed species.
Thus, it is challenging to predict and recommend the best among these three methods for all seaweeds.

Table 4 Nanocelluloses prepared from different seaweeds

S. Seaweed Product Treatments Crystalli- Size, References

N. Species nity Index Diameter
(%) (nm)
1 Cladophora Cellulose Mechanical 96.5 10-40 Xiang et
glomerata nanofibrils and al., 2016
enzymatic
2 Gelidium Nanocellulose Chemical 73 17.7-39.9 Chen et al.,
elegans treatments 2016
3 Laminaria Nanocrystalline Chemical 69.4 20-45 Liu et al.,
japonica waste cellulose treatments 2017a
4 Gelidiella Cellulose Mechanical 60 32 Singh et
acerosa nanocrystals and al., 2017
chemical
5 Red algae Cellulose Chemical 81-87 2.3-12.2 Achaby et
waste nanocrystals treatments al., 2018
6 Chaetomorpha Cellulose Chemical 85.02 30 Bhutiya et
antennina nanofiber treatment al., 2018
7 Sargassum Cellulose Mechanical 58-79 4.8-41 nm Doh et al.,
fluitans, nanocrystals and width, 21- 2020a
Laminaria chemical 248 nm
japonica, length

15
 Palmaria
palmate,
Porphyra
umbilicalis,
Ulva lactuca
8 Ulva lactuca cellulose Mechanical 63 -- Wahlstro¨
nanofibrils and m et al.,
chemical 2020
9 Laminaria cellulose Chemical 85-99 Length Doh et al.,
japonica, nanocrystals treatment 49-228, 2020b
Sargassum width 8-
natans 21 nm

To analysis, the suitability of nanocellulose prepared from seaweed-based cellulose for various
industrial applications, a comparison was made with the nanocelluloses produced from different
lignocellulosic biomass (Bamboo, Rice straw, and Eucalyptus) (Table 5). It was found that nanocelluloses
prepared from seaweed-based cellulose are higher crystallinity indexes (58-99%) and comparable sizes
(5-45 nm diameter) with those prepared from different lignocellulosic biomass (Table 5). Nanocellulose,
prepared from terrestrial biomass, is used widely in biomedical applications such as drug delivery, tissue
engineering, cardiovascular applications, wound dressings, cartilage replacements, and medical implants
(Guise & Fangueiro, 2016). Uses of nanocellulose-based polymer hybrids in biomedical engineering and
water purification are the emerging applications, where nanocellulose plays a vital role as a reinforcing
agent to improve materials' native properties (Patel et al., 2019). A few reports have investigated the
biomedical application of seaweed-based nanocellulose, especially from Cladophora (Bacakova et al.,
2019). Liu et al. (2017b) investigated the levels of various impurities present in Cladophora based
nanocellulose to test its applicability for biomedical applications and found it suitable for future
biomedical applications. Sulfonated Cladophora nanocellulose bead was prepared and tested for
hemocompatibility (Rocha et al., 2018) and its adsorption capacity for Congo Red dye (Ruan et al., 2018).
Apart from that, Cladophora based nanocellulose was also evaluated for its application as scaffolds for
cell cultivation (Metreveli et al., 2014). Other than Cladophora based nanocellulose, nanocellulose
prepared from Cystoseria myrica combined with Fe3O4 was studied to remove mercury ion pollution
(Zarei et al., 2018). However, there is more investigation needed to identify the potentials nanocellulose
from different algal species.

16
Table 5 Comparison of nanocelluloses prepared from seaweed-based cellulose and lignocellulosic
biomass

Biomass
Characteristics Seaweeds Rice straw Bamboo Eucalyptus
Crystallinity 58-99 62-76 52-70 34-60
Index (%)
Size in diameter 5-45 5-35 20-80 7.5-87
(nm)
References As per table Thakur et al., 2020; Lu et al., 2018; Hu Qing et al., 2013
4 Rezanezhad et al. et al., 2014
2013

5.2. Microcrystalline cellulose (MCC)

Microcrystalline cellulose, as revealed from its name, structurally, is long and thin fibers with a
three-dimensional network. Microcrystalline cellulose was discovered in 1955 by Battista and Smith and
commercialized under the brand name Avicel1 (Reier, 2013). Microcrystalline cellulose is produced from
cellulose with enzymatic or chemical pretreatment followed by mechanical treatment. Hydrochloric acid
is mostly used for the hydrolysis of cellulose. MCC is widely used in various industries as a stabilizer,
emulsifier, anticaking, and dispersing agent. The estimated global market value of microcrystalline
cellulose (MCC) was USD 885.1 million in 2018 and is expected to reach USD 1,241.4 million by 2023,
with a 7% compound annual growth rate (www.marketsandmarkets.com). Several reports emphasized the
preparation of microcrystalline cellulose from cellulosic material obtained from lignocellulosic biomass
(Beroual et al., 2021; Katakojwala & Mohan, 2020; Zhao et al., 2018; Thoorens et al., 2014). Recently, a
chemical–mechanical approach was used to prepare microcrystalline cellulose (MCC) from cellulose
extracted from brown seaweed Saccharina japonica (He et al., 2018). Cellulose was extracted from kelp
treated with hydrochloric acid and neutralized, followed by freeze-drying and grinding using a ball mill
with ZrO2 balls (1 cm diameter). MCC produced from Saccharina japonica showed high crystallinity
index (CrI) of 88.6%. The physical and chemical properties of prepared MCC were comparable with
commercial MCC prepared from a higher plant. However, more investigation is needed to explore the
suitability of the seaweed biomass to prepare microcrystalline cellulose for various commercial
applications.

17
5.3. Carboxymethylcellulose (CMC)

Carboxymethylcellulose (CMC) is prepared from cellulose by adding carboxymethyl groups (–

CH2- COOH) with some of the hydroxyl groups of the glucopyranose monomers of the cellulose
backbone (Heinze & Pfeiffer, 1999). The production of CMC comprises two steps, alkalization and
carboxymethylation, using chloroacetic acid serially (Lakshmi et al., 2017). CMC's physicochemical and
biological properties make it useful in various industrial sectors such as food, feed, pharmaceuticals,
agricultural and wastewater treatment (Ahmad et al., 2014; Borůvková & Wiener, 2013). Mainly, CMC
has been prepared from the cellulose extracted from terrestrial biomass. However, recently to find an
alternate renewable feedstock, a successful attempt was made to prepare CMC from cellulose isolated
from green seaweed Ulva fasciata (Lakshmi et al., 2017). The properties of synthesized CMC, like
molecular weight (MW) polydispersity and transparency, were comparable with commercial CMC.
Degree of Substitution (DS) 0.51 of algal cellulose-derived CMC was found higher than CMC from
Pomelo peel with 0.54 (Chumee & Seeburin, 2014) and sugarcane with 0.43 (Hong, 2013). Further, CMC
was used to prepare a film with a combination of Ag and leaf extract of Azadirachta indica and proved its
suitability for the film preparation. However, more detailed investigations from other seaweed species are
required to evaluate seaweed-based cellulose's potentials to prepare CMC and its commercial applications.

5.4. Paper preparation

The production of paper and paperboard is the primary non-food application of cellulosic pulp.
World production of paper and pulp was estimated at 419.72 million metric tons in 2018, and market value
projected to reach 523.6 billion U.S. dollars in 2022 (www.statista.com). Commercially cellulose is
obtained from wood pulp, and cotton is mainly used to manufacture paperboard and paper (Bogati, 2011).
Seaweed cellulose is projected as a raw material for paper-making as an alternative to wood-based
cellulose as seaweed can produce a larger mass of material in a shorter time than terrestrial-based plants.
It is also assumed that the use of seaweed cellulose for paper production will help overcome environmental
issues like deforestation and global warming (Mukherjee & Keshri, 2018).

Curtis et al. (1924), for the first time, patented the process for the preparation of pulp utilizing the
cellulose and alginate present in fresh alginophytic seaweed. Pulp comprises of the finely divided cellulose
having its fibrous structure substantially destroyed suspended in a solution of alginate and proposed its
utilization for binding or agglomerating agent. Further, a process was patented to prepare paper from
seaweed in combination with wood pulp (Nicolucci & Monegato, 1995). About 8-12 wt.% of seaweed
particles (≤500 µm) prepared from the green seaweed Ulva rigida and Ulva lactuca mixed with bleached
wood-pulp and calcium carbonate fed into a paper-making machine. The use of algal material improved

18
the mechanical (resistance to bursting, stiffness, and rupture length) and chemical characteristics of the
paper. The paper was recently prepared using bleached pulp produced from two red algae species,
Gelidium corneum, Gelidium amansii, and paper properties compared with bleached wood chemical
pulps-based paper (Seo et al., 2010b). The paper prepared from red algae pulp was found superior to wood
pulp-based paper in high Bekk smoothness and opacity, which are essential properties for quality printing
paper. The commercial production of quality papers from macroalgal cellulose has a long way to go, and
it needs a high volume of cellulose and extensive investigation.

5.5. Bioplastic

Polymers possess a wide range of properties and became a part of everyday life (Cheng, 2008).
The polymers are classified into two categories -synthetic and biopolymers. Plastic is one of the best
examples of synthetic polymers which have an important role in everyday human life globally. Synthetic
polymers are made up of carbon-carbon bonds as their backbone, typically derivatized in a controlled
environment from petroleum oil (Shrivastava, 2018). Properties of synthetic polymers such as strength,
high elasticity, resistance to chemical, and physical degradation make them unique and useful (Bhadra et
al., 2014). However, these polymers' non-degradable nature has become a severe environmental crisis
(Kyrikou & Briassoulis, 2007). The disposal of synthetic plastic waste becomes a major challenge for the
world, and now attention is turning towards biopolymers. The properties like renewability,
biodegradability, and biocompatibility are the major characteristics of biopolymers-based plastic making
it suitable for several industrial and medical applications (Dassanayake et al., 2018). However, bioplastics'
high water absorption tendency is one of the potential drawbacks as it may alter the resulting plastic's
elastic modulus, thus making plastic with a lower elasticity. Similarly, the low thermal stability of
bioplastics limits its applicability.

Production of bioplastic is one of the emerging applications of seaweeds and their cellulose. The
bioplastics were synthesized using sorghum and red seaweed Eucheuma spinosum fiber (Darni et al.,
2019). The properties of prepared bioplastic such as tensile strength (21.265 MPa), elongation (4.467%),
Young’s Modulus (498.463 MPa), density (0.95 g/cm3), and water uptake (21.265%) was comparable
with characteristics of commercial HDPE plastic. Recently, nanocomposite films were prepared using
crude extracts and cellulose nanocrystals (CNCs) isolated from two seaweeds, Laminaria japonica, and
Sargassum natans, with suggested applications in food preservation or to design functional foods (Doh,
2020c). The uses of cellulose nanocrystals (CNCs) improved the physicochemical, mechanical, barrier,
and thermal properties of biofilms. Similarly, Kamarol Arifin (2020) developed biofilm using a
combination of cellulose extracted from a green seaweed Caulerpa lentillifera and corn starch that showed
good biodegradable properties. The bioplastic film prepared with 10% cellulose concentration showed the

19
highest tensile strength and increased Young Modulus value. Overall, studies on the uses of seaweed-
based cellulose to prepare bioplastics confirmed that it could be used as a filler in producing bioplastics.

5.6. Production of cellulosic ethanol

The non-renewable nature of petroleum-based fuels and the negative consequences on the
environment of their usage has given rise to find an appropriate and efficient alternative. Among the
possible alternative to petroleum fuels, bioethanol production is the primary target globally. Most ethanol
produced is first-generation bioethanol which comes from plant-derived sugar such as sugarcane, sugar
beet, corn, and wheat. Due to the rising concern over the “Food vs.Fuel”; the lignocellulosic materials for
bioethanol production got highlighted (Baghel et al., 2015). However, removing lignin from such biomass
before hydrolysis is a major challenge affecting the economics of bioethanol production and generating
chemical effluents (Trivedi et al., 2013). After first and second-generation resources, seaweed biomass
emerged as a third-generation resource for bioethanol production (Kumar et al., 2013).

Seaweeds constitute high carbohydrate, low lignin content, and a higher productivity without any
agricultural inputs. Firstly, bioethanol was produced from the extract of brown seaweed Laminaria
hyperborean (Horn et al., 2000). Afterward, various researchers reported the potential of different
seaweeds for bioethanol production (Lee & Lee, 2012; Ge et al., 2011; Adams et al., 2009; Uchida &
Murata, 2004). Although there were many developments in seaweed-based bioethanol research, the
challenge remains to achieve high ethanol yields. The chronology for the production of bioethanol from
seaweed biomass is presented in Figure 5. As only cellulose fraction of seaweed biomass, which is in
minor proportion, is easily hydrolyzed and fermented while a larger part of pentose type of sugar is
difficult to hydrolysis and ferment using common routes. Later, an engineered microbial platform for
direct bioethanol production using either alginate alone (Takeda et al., 2011) or whole biomass (Enquist-
Newman et al., 2014; Wargackiet al., 2012) has also been developed to utilize the maximum carbohydrates
fraction. However, a concern was developed that utilization of whole polysaccharides of seaweed whole
polysaccharides for bioethanol production could not sustain for a long time (Baghel et al., 2015).
Polysaccharides such as agar, carrageenan, and alginate extracted from seaweed biomass are high-value
products compared to bioethanol and have established a global market. The commercial uses of such
polysaccharides fraction for bioethanol in the long term may jeopardize the multibillion seaweed
hydrocolloid industries. Such seaweed biomass could be utilized for bioethanol production in the case of
surplus biomass production only. Subsequently, the attention has been shifted towards the production of
ethanol from only seaweed cellulose fraction in biorefinery approaches. The extraction of cellulose from
3 different agar-producing seaweeds after recovery of pigments, mineral, lipid, and agar was reported, and
further cellulose was hydrolyzed and fermented to produce bioethanol (Baghel et al., 2015). Similarly,

20
bioethanol was produced from cellulose present in residual waste of green seaweed Ulva fasciata and red
seaweed Gracilaria corticata (Trivedi et al., 2016; Baghel et al., 2016).

Figure 5 Strategies for bioethanol production from seaweeds

6. Conclusion and future perspectives

Cellulose extracted from various seaweed species has shown similar properties to those produced from
lignocellulosic resources. The lower content of cellulose in seaweed biomass is a major drawback;
negligible or no lignin content is beneficial. The seaweed cellulose-based products could be used in
various industrial applications, as proved in various research reports. There is an immense opportunity to
extract seaweed cellulose and their application development with the growing seaweed global production.
Based on a detailed analysis of various research papers published on seaweed cellulose, it can be inferred
that (1) The yields of cellulose for most seaweed species are not comparable with those from
lignocellulosic biomass. (2) The cellulose yields (27-45%) from residual seaweed waste are comparable
with that of lignocellulosic biomass and need efforts to utilize such waste to produce cellulose and create
value from the waste. (3) It could be feasible and economical to extract cellulose from seaweed biomass
as a co-product. (4) There is a need for technological inventions for large-scale farming of cellulose-rich
algal species such as Cladophora, Ulva, Gelidiella, and Gelidium for cellulose production. (5) Since
demand increases for cellulose-based soft materials, using seaweed-based cellulose to prepare high-value
products such as nanocellulose and their application as soft material in food and cosmetics could be a
topic of future research.

21
Acknowledgement

The financial support received from OLP-2005 is gratefully acknowledged. RSB thankful to the Director,
CSIR-NIO (CSIR-National Institute of Oceanography) for extending the facilities.This is CSIR-NIO’s
contribution reference number XXXX. RPS would like to thanks toThe Department of Biotechnology for
providing Ramalingaswami-re-entry fellowship.

22
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