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Fucoidans from brown seaweeds: an update on structures, extraction techniques and use of enzymes as tools for structural elucidation
Open accessAbstract
Fucoidan or fucoidans cover a family of sulfated fucose-rich polysaccharides, built of a backbone of L-fucose units, and characteristically found in brown seaweeds. Fucoidans have potential therapeutic properties, including anti-inflammatory and anti-coagulant activities, as well as anti-proliferative effects on cancer cells. Recent work has revealed distinct structural features of fucoidans obtained from different brown seaweed sources. Fucoidans are classically obtained from brown seaweeds by multi-step, hot acid extraction, but the structural and compositional traits, and possibly the bioactivity, of the fucoidan polysaccharides are significantly influenced by the extraction parameters. This review discusses the structural features of fucoidans, the significance of different extraction technologies, and reviews enzymatic degradation of fucoidans and the use of fucoidan-modifying enzymes for elucidating structural details of fucoidans. Mild extraction techniques coupled with the use of new tools such as enzymes can provide the much needed knowledge about structural characteristics of different fucoidan molecules and thus pave the way for a better understanding of the structural chemistry and bioactivities of fucoidans.
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TL;DR
This review synthesizes the structural diversity of fucoidans (fucose-containing sulfated polysaccharides, FCSPs) from brown seaweeds, arguing that extraction conditions strongly shape yield and composition and that no consistent "fucoidan" backbone exists. It also reviews fucoidan-modifying enzymes (fucoidanases, fucosidases) as tools for structural elucidation and discusses bioactivity and commercial prospects.
Key findings
- Fucoidan structures vary by species and cannot be predicted by algal order; the prevalent core backbones are (1→3)-linked α-L-fucopyranose and alternating (1→3)/(1→4)-linked α-L-fucopyranosyls, with variable sulfate, acetate, and branch substitutions (Table 1).
- Extraction parameters significantly affect yield and composition: in the authors' own single-step study on *Sargassum henslowianum*, temperature and time had positive effects and acid concentration a negative effect, with a maximal fucoidan yield of ~7% at 0.03 M HCl, 90 °C, 4 h; 0.2 M HCl broke polysaccharide integrity, lowering fucose at elevated time/temperature.
- Historical extraction data: a single acid hydrolysis recovered ~50% (w/w) of theoretical maximum fucoidan, while three rounds recovered >80% of fucose; the triple-step protocol (0.1 M HCl, pH 2.0–2.5, 70 °C, 1 h ×3) gave crude fucoidans with 30–36% fucose, and *F. vesiculosus* product contained 44% fucose, 26% total sulfate, and 31% ash. Yields as % fucose of total seaweed fucose were *Pelvetia canaliculata* 76%, *F. vesiculosus* 62%, *Ascophyllum nodosum* 54%, *L. cloustoni* 20%.
- Fucoidanases show specificity: an endo-1,3-α-fucoidanase from *Fucanobacter lyticus* degraded Laminariales fucoidans but not Fucales or Chordariales; enzymatic treatment of *Fucus distichus* fucoidan confirmed repeating disaccharide units [→3)-α-L-Fucp-(2,4-di-SO₃²⁻)-(1→4)-α-L-Fucp-(2-SO₃²⁻)-(1→].
Methods
- Narrative review covering fucoidan structural chemistry, extraction technology history (1913–1950s onward), extraction-parameter effects, and fucoidan-modifying enzymes.
- Includes the authors' own single-step extraction experiment using *Sargassum henslowianum* as a model brown seaweed, with HPAEC-PAD analysis of TFA hydrolysates; extraction parameters tested were acid concentration, time, and temperature.
- Enzyme data are compiled from published studies (Table 4) and classified by cleavage pattern: type 1 (1→4)-α-L-fucoidan endohydrolase, type 2 (1→3)-α-L-fucoidan endohydrolase, and type 3 exo-acting α-L-fucosidase.
Limitations
- Stated: no standardized purification procedure exists for fucoidans; the use of many different extraction techniques has exacerbated structural elucidation; the exact structural elements responsible for bioactivity remain unidentified; no pharmaceutical, dermatological, or nutraceutical applications have been officially approved.
- Stated: heterogeneity within the same seaweed species can lead to pronounced differences in extracted products.
- Evident from the text: as a narrative review it does not describe a systematic literature search or inclusion criteria, and the extraction-parameter findings derive largely from a single model species (*S. henslowianum*), limiting generalizability.
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48 Not checked yet · 21 Own interpretation · 19 No basis given · 3 Own result · 1 Supported so far
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In the paper
“Structural integrity of the polysaccharide was best conserved at low acid treatment.”
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In the paper
“A maximal fucoidan yield of y7% was achieved by the optimal extraction procedure of: 0.03 M HCl, 90 uC, 4 h.”
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In the paper
“The HPAEC-PAD analysis of the TFA hydrolysate of S. henslowianum confirmed that fucose, galactose, and glucuronic acid were the major constituents of the extracted polysaccharides obtained by the optimized method.3”
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- Not checked yetcites [3] source
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In the paper
“The use of 0.2 M HCl apparently broke the integrity of the polysaccharide molecules resulting in a decline of fucose at elevated time and temperature.3”
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In the paper
“the structural and compositional traits, and possibly the bioactivity, of the fucoidan polysaccharides are significantly influenced by the extraction parameters.”
- Also stated in: § 3.1 Influence of extraction parameters on fucoidan yield and composition · boosted
“The data demonstrate the vulnerability of fucoidan structures to harsh extraction conditions and confirm that the extraction method significantly influences the yields and not least the composition of the extracted polysaccharides.”
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- Also stated in: § 3.1 Influence of extraction parameters on fucoidan yield and composition · boosted
In the paper
“Several of the fucoidan structures reported in the literature have also contained small amounts of various other monosaccharides e.g.glucose, galactose, xylose, and/or mannose. 1”
What the sources say
- Not checked yetcites [1] source
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In the paper
“The differences caused by differences in extraction methodologies obviously may have major bearings on the results when the fucoidan polysaccharide products are being evaluated for biological activity.”
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In the paper
“Likewise, the enzymatic treatment confirmed that the main fucoidan structural unit from this brown seaweed Fucus spp.consists of repeating di-saccharide units of [A3)-a-L-Fucp-(2,4di-SO 3 2 )-(1A4)-a-L-Fucp-(2SO 3 2 )-(1A].35”
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- Not checked yetcites [35] source
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In the paper
“The variations in the reported structural properties of fucoidans from different brown seaweed species clearly confirm the natural biodiversity of fucoidans as exemplified by the structures found in different seaweed taxonomical orders, notably Fucales, Laminariales and Chordariales (Table 1).”
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In the paper
“The bioactivities of fucoidans are related to their structural make-up, monosaccharide composition, sulfate content, position of sulfate ester groups, and molecular weight.2”
What the sources say
- Not checked yetcites [2] source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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- Rests on an assumption: Fucoidan polysaccharides have potential therapeutic properties, including anti-inflammatory, anti-coagulant, and anti-proliferative effects on cancer cells.
- It may state two separate claims as one. 33
- Probably not a substantial claim. 14
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In the paper
“Algal fucoidans have been the subject of numerous scientific studies due to their diverse biological functions and possible therapeutic properties including antitumor, immunomodulatory, 45 anti-inflammatory, 6 antithrombotic, and anticoagulant effects.46”
What the sources say
- Not checked yetcites [45] source
- Supportedcites [6] source
Citation 46 is attached to the antithrombotic and anticoagulant effects, not the earlier effects, which have separate citations. The Discussion reports prolonged clotting time for most fucoidans and strong inhibition of thrombin-induced platelet aggregation by five fucoidans; the Introduction describes their potential as drugs for thrombosis. (LLM, conf=0.92)
- Not checked yetcites [46] source
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In the paper
“All extraction parameters had significant effects on the yields, i.e. the fucose-containing sulfated polysaccharide yield, with the temperature and time exerting positive effects, and the acid concentration having a negative effect.”
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In the paper
“Lower polysaccharide yield, but relatively higher fucose content was obtained with shorter extraction time.”
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In the paper
“Fucoidan or fucoidans cover a family of sulfated fucose-rich polysaccharides, built of a backbone of Lfucose units, and characteristically found in brown seaweeds.”
- Also stated in: § 1 Introduction · plain
“Fucoidans in fact designate a group of fucose-rich, sulfated polysaccharides, usually built of a backbone of a-linked L- fucose residues having various substitutions.”
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- Not checked yetcites [1] source
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- Also stated in: § 1 Introduction · plain
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In the paper
“Enzymes catalyzing modifications of fucoidan polysaccharides, i.e. ''fucoidanases'', include: i) Fucoidanases, EC 3.2.1.44,which catalyze the endo-hydrolysis of (1A2)-a-L-fucoside linkages in fucoidans without release of sulfate, and ii) a-L-Fucosidases, EC 3.2.1.51which catalyze the hydrolytic release of L-fucose, via cleavage of a(1A3)-, a(1A4)and possibly also of a(1A2)-O-fucosyl bonds in fuc”
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In the paper
“It has also been shown by others that longer extraction time at higher temperatures led to higher polysaccharide yield with lower amounts of sulfate and a higher proportion of glucuronic acid.11,17”
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In the paper
“The data demonstrate the vulnerability of fucoidan structures to harsh extraction conditions and confirm that the extraction method significantly influences the yields and not least the composition of the extracted polysaccharides.”
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In the paper
“Fucoidan polysaccharides are mainly found in brown seaweeds, but fucoidans differ in structure among brown seaweed species.”
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“Fucoidans isolated from L. saccharina (Laminariales) are thus mainly composed of a(1A3)-linked L- fucopyranose units, but with sulfate at both C-2 and C-4, or at C-4 alone, however (Table 1).”
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In the paper
“These fucoidan molecules have a backbone consisting of a repeating structure of a(1A3)-linked L-fucopyranose residues with sulfate at the C-2 position, with the a(1A3)-linked L- fucopyranose residues linked to a(1A4)-L-fucopyranose residues with disulfate at C-2 and C-3 (Table 1).”
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In the paper
“The sulfate groups on the fucans are typically located at C-4 of a (1A3)-linked unit or on C-2 of a (1A4)-linked residue.11”
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In the paper
“and may include calcium addition to promote alginate precipitation during purification, 3,4”
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In the paper
“It has recently been demonstrated that fucoidans are vulnerable to harsh extraction conditions and that the extraction method significantly influences the yields and not least the polysaccharide composition of the extracted fucose-containing sulfated polysaccharides.3”
What the sources say
- Not checked yetcites [3] source
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In the paper
“On the other hand, fucoidanases have been intensely studied in relation to bacterial degradation of fucoidans and molecular structure elucidation of fucoidan”
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In the paper
“Mild extraction techniques coupled with the use of new tools such as enzymes can provide the much needed knowledge about structural characteristics of different fucoidan molecules and thus pave the way for a better understanding of the structural chemistry and bioactivities of fucoidans.”
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In the paper
“Similarly, the fucoidans from Cladosiphon okamuranus (Chordariales) are basically made up of linear backbones of a(1A3)-linked L- fucopyranose residues (Table 1) with a portion of the fucose residues being O-acetylated and with some sulfate substitution at the C-4 position.8”
What the sources say
- Not checked yetcites [8] source
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In the paper
“hand, fucoidanases have been intensely studied in relation to bacterial degradation of fucoidans and molecular structure elucidation of fucoidan polysaccharides.35,36,42”
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In the paper
“These structural units are similar to those found in the fucoidans from Chorda filum (Laminariales) which consist of a core backbone structure of poly-a(1A3)-linked L-fucopyranoses sulfated mainly at C-4, sometimes at the C-2 position, and with some of the a(1A3)linked fucose residues being 2-O-acetylated.4,14”
What the sources say
- Not checked yetcites [4] source
- Not checked yetcites [14]
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In the paper
“Sulfated b-D-galactan has been isolated in trace amounts from Laminaria angustata and these polysaccharides have also been classified as fucoidans, eventhough b-D-galactose appear to be the major sugar, and L-fucose and D-glucuronic acid are present only as trace components.15”
What the sources say
- Not checked yetcites [15] source
hedged- Asserted: passes — The authors put it forward as true.
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In the paper
“The fucoidans in brown seaweeds in the order of Fucales, e.g.Fucus evanescens and Fucus serratus, represent the typical structures: They possess a large proportion of both a(1A3)and a(1A4)-linked L-fucopyranose residues, which may be substituted with sulfate (SO 3 2 ) on C-2 and C-4 (Table 1).6,7,10”
What the sources say
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In the paper
“It also seems clear that fucoidans extracted from brown seaweeds under the same seaweed classification (i.e.order) have different composition, and in turn that the structural traits cannot be categorized or predicted according to algal order as also indicated in Table 2.”
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In the paper
“Mild extraction technology is essential for maintaining the structural integrity of the fucoidan polysaccharides and for preserving their biological properties.”
plaindirect- Asserted: passes — The authors put it forward as true.
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- Rests on an assumption: Fucoidan polysaccharides have potential therapeutic properties, including anti-inflammatory, anti-coagulant, and anti-proliferative effects on cancer cells.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The available published data, including our own findings, thus generally show that the term ''fucoidan'' has been used for several chemical structures, and vice versa, that fucoidans is a term that covers a family of fucose-containing sulfated polysaccharides.”
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In the paper
“Ponce et al. (2003) 17 thus reported that longer extraction time led to poorer fucose content, whereas glucuronic acid increased while sulfate decreased with time, i.e., the sulfate content was highest when glucuronic acid was lowest.17”
What the sources say
- Not checked yetcites [17] source
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In the paper
“Fucoidans from brown seaweed algae generally consist of a backbone of a(1A3)-L-fucopyranose residues or of alternating a(1A3) and a(1A4)-linked L-fucopyranosyls, which in either case may be substituted with sulfate or acetate and/or have side branches containing fucopyranoses or other glycosyl units, e.g.glucuronic acid.”
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In the paper
“Since fucoidans are water soluble, they can easily be incorporated into lotions, creams and other beauty products.”
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In the paper
“Marine bacteria such as Fucophilus fucoidanolyticus are to a certain extent able to consume fucoidans of various structures and apparently”
What the sources say
- Not checked yetcites [44] source
hedgedconditional- Asserted: passes — The authors put it forward as true.
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In the paper
“In this connection, the targeted use of fucoidanases might assist in determining the precise molecular structures accountable for preventive bioactivity effects.”
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In the paper
“Fucoidans may enhance dermal fibroblast proliferation and deposition of collagen; 52”
What the sources say
- Not checked yetcites [52] source
hedgedconditional- Asserted: passes — The authors put it forward as true.
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In the paper
“Production of pure fucoidan products is challenging because of the presence of other polysaccharides, such as alginate.”
plaindirect- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Among the polysaccharides from brown seaweeds, fucoidans have turned out to exert potentially significant pharmaceutical effects, but at the same time to be a family of differently structured polysaccharides comprising structural units having distinct traits.”
hedged- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“It is known now that the fine structural details of fucoidans vary depending on seaweed type and species.”
boostedcorrelational- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper may give no reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The lack of a standardized extraction methodology prevents detailed comparison and most importantly obstructs the understanding of the correlation between fucoidan 4.”
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
In the paper
“At least one of them (intracellular fucoidanase) seems to have the ability to catalyse the splitting of 1,3a-glycosidic bonds and has also been demonstrated to express deacetylase activity being effective on acetylated fucoidans from Cladosiphon okamuranus.33”
What the sources say
- Not checked yetcites [33] source
hedgedconditional- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“The biological activity of fucoidans depend on the seaweed source, the compositional and structural traits, the content (charge density), distribution, and bonding of the sulfate substitutions, and the purity of the fucoidan product.”
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
In the paper
“It has been suggested that the bioactive properties of fucoidans are defined by their structural make-up, and notably related to the fucose sulfated chains.15”
What the sources say
- Not checked yetcites [15] source
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Despite intensive research, the exact correlations between the various bioactivities and the molecular and structural features of fucoidans still have to be clarified.”
plaincorrelational- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim.
None of the sources cited for this claim have been checked yet.
In the paper
“Fucoidan was also reported to induce apoptosis in HT-29 colon cancer cells, 47 MCF-7 human breast cancer cells, 48 and HS-Sultan human lymphoma cells.49”
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
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It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“As mentioned earlier, fucoidan polysaccharides and other FCSPs exhibit significant biological properties in vitro such as anti-prolif-erative activity on cancer cells.”
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. The paper may give no reason why it holds.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Harsh chemicals are environmentally burdensome and have detrimental effects on the yield and compositions of the end product polysaccharides.”
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
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Probably not a substantial claim. The paper may give no reason why it holds. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Fucoidanases may provide new tools for elucidation of detailed fucoidan structures from various sources.”
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: fails — It may be too vague to check against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. It may be too vague to check against evidence. The paper may give no reason why it holds.
The paper relies on this without support.
Fucoidan polysaccharides have potential therapeutic properties, including anti-inflammatory, anti-coagulant, and anti-proliferative effects on cancer cells.
“Fucoidans have potential therapeutic properties, including anti-inflammatory and anti-coagulant activities, as well as anti-proliferative effects on cancer cells.”
“Fucoidan polysaccharides exhibit a number of pharmaceutically interesting biological activities, including immuno-modulatory, anti-tumor, anti-inflammatory and anti-coagulant effects (to be discussed in more detail later).”
The paper relies on this without support.
No standardized purification procedure is available for fucoidans at present.
“but no standardized purification procedure is available for fucoidans at present.”
The paper relies on this without support.
The heterogeneity of the structural make-up of seaweeds within the same species may lead to pronounced differences in the products extracted.
“In addition, the heterogeneity of the structural make-up of seaweeds within the same species may lead to pronounced differences in the products extracted.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
In the citing paper
“The lack of standardized extraction methodologies has prevented the approval of polysaccharides or their derived fractions to be officially approved for pharmaceutical, dermatological, nutraceutical or other commercial applications to date”
From Polysaccharides from macroalgae: Recent advances, innovative technologies and challenges in extraction and purification (2017) · cites this paper as Ale & Meyer, 2013
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
Rated minor: it would not touch a result — a value no result rests on, a p-value that stays on the same side of the significance level, or a conflict that is clerical, resolved elsewhere in the paper, or outside its own findings.
In the paper
§ Laminaria religiosa
“By this procedure, samples of crude fucoidans containing 30-36% fucose were obtained; for example, the fucoidan polysaccharides recovered from F. vesiculosus using this optimal extraction protocol was analyzed to contain (by weight) 44% fucose;”
§ Laminaria religiosa
“samples of crude fucoidans containing 30-36% fucose were obtained”
Citations
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Metadata
- Authors
- Ale, M. T., & Meyer, A. S.
- Affiliations
- Technical University of Denmark (DK)
- Corresponding author
- Meyer, Anne S.
- Venue
- RSC Adv.
- Publisher
- Royal Society of Chemistry (RSC)
- Published
- 2013
- Volume
- 3
- Issue
- 22
- Pages
- 8131-8141
- ISSN
- 2046-2069 (ISSN-L)
- OpenAlex
- W2165865189
- Semantic Scholar
- 9ec7156ada540774a377f33054ce052291b5f9dc
- Cited by
- 381
- References
- 50
- Citations by year
- 2026: 38 · 2025: 33 · 2024: 32 · 2023: 33 · 2022: 36 · 2021: 39 · 2020: 50 · 2019: 43 · 2018: 28 · 2017: 17 · 2016: 13 · 2015: 9 · 2014: 9
- FWCI
- 15.78
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- cc-by
- Locations
- RSC Advances (publishedVersion, cc-by)
- Technical University of Denmark, DTU Orbit (Technical University of Denmark, DTU) (submittedVersion, cc-by)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Aquatic Science › Seaweed-derived Bioactive Compounds
- Life Sciences › Agricultural and Biological Sciences › Food Science › Polysaccharides Composition and Applications
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Polysaccharides and Plant Cell Walls
- Keywords
- fucoidan, brown algae, polysaccharide structure, enzymatic degradation, structure elucidation, anti-inflammatory activity, anticoagulant activity, antiproliferative activity
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Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
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