Advanced Composites and Hybrid Materials (2024) 7:27

https://doi.org/10.1007/s42114-024-00836-3

REVIEW

Transforming lignin into value‑added products: Perspectives on lignin

chemistry, lignin‑based biocomposites, and pathways for augmenting
ligninolytic enzyme production
Subhashree Rath1 · Deepak Pradhan2 · Haishun Du3 · Sonali Mohapatra3,4 · Hrudayanath Thatoi1,4

Received: 21 April 2023 / Revised: 7 November 2023 / Accepted: 12 January 2024 / Published online: 7 February 2024
© The Author(s), under exclusive licence to Springer Nature Switzerland AG 2024

Abstract
Lignin is a promising biopolymer abundantly used in industrially valued biobased products. But the resilient nature of lignin
reinforces difficulties during its depolymerization, leading to distinct challenges in manufacturing value-added products.
This review aims to provide information on lignin chemistry that imparts thoughtful insights on development of lignin-based
biocomposites and hybrid materials that are both economical and environment convivial products. Biological approaches
on ligninolytic microorganisms that are also attractive pathways for conversion of lignin into value-added products are
elaborated. Further, this review also emphasizes on the recent metagenomics, system biology, and in silico approaches on
enhancing the ligninase production from microbial sources and their utilization for the synthesis of lignin-based materials.
Finally, updates on the patented lignin biocomposites and hybrid materials have been showcased. It is expected that the
review will address the present necessity of waste management issues associated with agro-wastes and simultaneously provide
understanding for the technology-based approaches that can be utilized for manufacturing feasible industrially value-added
products from lignin as a biomass.

Keywords Lignin hybrid materials · Lignin biocomposites · Lignolytic enzymes · In silico · Metagenomics

1 Introduction products such as composites and hybrid materials has been an

Sustainable development for improving human society and the
environment is an integral part that has feast into all fields
including industrial sectors that promotes development of bio-
logical product [1]. Developing enzymes that transform agro-
wastes from renewable lignocellulosic biomass to value-added
* Sonali Mohapatra
smohapatra4@wisc.edu
* Hrudayanath Thatoi
hnthatoi@gmail.com
1 polypropylene composite, lignin-epoxy composite resin, lignin
Department of Biotechnology, Maharaja Sriram Chandra Bhanja
Deo University, Takatpur, Baripada 757003, Odisha, India
2 hybrid materials, etc.) appears to be a rational approach from
School of Pharmaceutical Sciences, Siksha ‘O’ Anusandhan
(Deemed to Be University), Bhubaneswar 751003, Odisha,
India
3
Dept. of Biological Systems Engg, University of Wisconsin–
Madison, 1710 University Avenue, Madison, WI 75376, USA
4
Present Address: Center for Industrial Biotechnology,
Sikha ‘O’ Anusandhan Deemed to be University,
Bhubanseswar 751003, Odisha, India
area of interest in the recent years [2]. Lignin, an aromatic pol-
ymer with a benzene ring, is the second most abundant (consti-
tutes about 30%) cheap lignocellulosic recourse that provides
rigidity to cell walls and pockets the cellulose and hemicellu-
lose in vascular plants. Approximately 50 million tons of lignin
is extracted annually from the pulp and paper industries and
thirty million tons from ethanol and textile industries as by-
products [3]. From these by-products, < 2% of lignin has been
commercialized to date, and the major portions are largely
burnt, generating deep environmental concerns. In this sce-
nario, the conversion of lignin to value-added products (lignin-
rubber composite, lignin polysaccharide composites, lignin
an economic and ecological perspective. Moreover, wood-
based or lignin composite materials can offer efficient alter-
natives to materials like traditional petroleum-derived plastics,
contributing to the reduction of greenhouse gas emissions [4,
5]. Lignin-based materials are typically eco-friendly and cost-
effective, finding extensive applications in energy storage,
environmental solutions, electronic devices, and various other

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27 Page 2 of 29
industries [6]. However, modification of recalcitrant lignin that
consists of a densely packed three-dimensional network of phe-
nylpropanoid building blocks, i.e., coniferyl, coumaryl, and
sinapyl alcohol in different ratios to high-value-added materi-
als and chemicals, is challenging due to their distinctive chemi-
cal structure and reactivity nature [7]. Lignin conversion can be
conducted chemically, physically, or biologically. The dense,
complex structures of lignin bounded by ether linkages and
carbon-carbon bonds are very strong, and it is difficult to break
them by only using chemicals Moreover, the high temperatures
(up to 500°) and high pressures (up to 200 bar) required by the
physical methods for lignin breakdown significantly increase
the cost and quality of the lignin [8]. Biological methods by
use of bacteria, actinomycetes, fungi, and yeasts although are
economical, but are slow process mostly because of the com-
plexity and low nitrogen content of lignin as a substrate [9]. In
this context, manipulating the microbes for the production of
higher titers of these driver enzymes using lignin as a substrate
is an area of recent interest to the scientific community. How-
ever, considerable knowledge of the interactions of enzyme
production system of lignolytic microorganisms with other
lignin-degrading communities and acquaintance of the pro-
duction routes of these enzymes can be beneficial in designing
industry-efficient microorganisms.
Metagenomic profiling, synthetic biology approach, and
in silico studies are accommodating tools in determining
the aforesaid prospects. Metagenomic profiling can help
target the vital relationship between the lignin-degrading
microbes in the environment or their living counterparts
with the microbial community of other conglomerates.
Scully et al. [10] investigated the large-scale comparative
study between the mid-gut metagenome of Anoplophora
glabripennis (wood-feeding beetles) and their microbial
associates that add to the breakdown of woody tissue. The
study identified four essential lignin-degrading enzymes,
thirty-six families of glycoside hydrolases, and nutrient
recovery genes that could serve as a pool of novel enzymes
for industrial lignin utilization. Likewise, synthetic biology
approaches that employ fermentative recombinant microbial
approach are more dominant than genetic engineering routes
[11]. Synthetic routes deploy optimization of distinct parts of
the defined enzyme pathway, such as the enzyme homologs,
promoters, and ribosomal binding sites, to enhance the effi-
cacy of the pathway [12]. However, metagenomic and syn-
thetic routes without an apposite assessment may produce
mispredictions, specifically in genes with numerous exons
and very short intron/exon sizes [13]. Hence, prior in-silico
studies can provide valuable insights into predicting the
most appropriate pathway to improve the metagenomic and
synthetic processes further.
Thus, this review aims to describe the different charac-
teristics of lignin’s chemical structure from various sources,
the primary enzymes involved in the ligninolysis of lignin,
Advanced Composites and Hybrid Materials (2024) 7:27
and the utilization of these enzymes for the degradation of
lignin in the production of chemical and lignin composite
materials. The study elaborates a thorough assessment of
the metagenomic, systems biology, and in silico–based
approaches for enhancing ligninolytic enzyme production
and their utilization for the synthesis of lignin-based materi-
als. Finally, it apprises on the patented lignin biocomposites
and hybrid materials have been showcased.
2 Characteristics of lignin from different sources
Lignin is a heterogeneous biopolymer that is hydropho-
bic and is vital for conferring structural stiffness to plants,
effective transport of water systems in the plant, and non-
invasiveness of external elements through the conduits.
Lignin is predominantly found in the vessels and fibers of
the plant cell wall entwined with hemicellulose and cel-
lulose microfibrils through chemical bonds. Apart from
providing stability, it also plays a significant role in pro-
tecting plants during infection by impeding the entry of the
cellulolytic enzymes from bacteria and fungus and subse-
quently obstructing pathogen invasion to the surrounding
tissues. The customary lignin structure in the lignocellulose
is formed by block co-polymerization in the environment,
i.e., an integrated enzymatic dehydrogenate polymerization
of three phenylpropanoid units, coniferyl alcohol, synapyl
alcohol, and p-coumaryl alcohol. Based on methoxy substi-
tution on the aromatic ring of phenylpropanoid, the major
backbone units in p-courmaryl alcohol are p-hydroxyphenyl
(H); in coniferyl alcohol, it is guaiacyl (G); and in sinapyl
alcohol, it is syringyl (S) [14] (Fig. 1).
The H/G, S/G, and H/S/G ratio that is reliant on the plant
growth stage and environmental factors are the major con-
tributors that elect the fate of lignin composition and its
content in lignocellulosic biomass between different species
and within the components of the same species (i.e., lignin
content in wood and the bark). Further, β-O-4 linkage that
serves as the backbone for the muster of H, S, and G units;
the phenyl coumarins; and the resinols are also determinant
factors for lignin content in biomass [15]. Thus, the consid-
erate approach to utilizing lignin requires a thorough under-
standing of lignin chemistry specific to different biomass.
The broad classification for different lignin sources
can be based on the primary division of plant biota, i.e.,
gymnosperms, angiosperm dicotyledons, and angiosperm
monocotyledons. The gymnosperms mostly deposit lignin
that is derived primarily from G monomers together with a
small proportion of p-hydroxyphenyl units. Gymnosperm
lignins generally have fewer beta-O-4 ether bonds, higher
phenolic hydroxyls, and greater cross-links. Addition-
ally, the subunits are allied by α-O-4 linkages, β-β link-
ages, β-5 linkages, 5–5 linkages, and biphenyl and diaryl
Advanced Composites and Hybrid Materials (2024) 7:27
Fig. 1  Lignin compounds with different functional groups and interlinkage
ether structures (Fig. 1). In angiosperm dicotyledon,
lignins are composed of G and S units, whereas lignin
of angiosperm monocotyledons contains p-coumaryl
alcohol units (G/S/H-type lignin) [16]. Interestingly,
the dissemination and prevalence of the different
monolignols are critical elements for the amount and
nature of the interknit linkages. For instance, monolignols
with ortho-methoxy-groups cannot form 5–5 and β-5
carbon-carbon bonds which is a characteristic feature of
angiosperm monocotyledons, whereas the gymnosperm
lignins consist of two methoxy-substituted ortho-groups.
Del Río et al. [17] have demonstrated a perfect picture
of the structural difference between angiosperms and
gymnosperms. The differences are favorably over S units,
wherein the angiosperms are extensively acylated at the
γ-carbon of the lignin side chain (up to 80% acylation)
Page 3 of 29 27
with acetate and/or p-coumarate groups, whereas the
gymnosperms are esterified with acetate and p-coumarate
groups. Consequently, the processing of these angiosperms
and gymnosperm biomass through different pretreatment
strategies (milling, acid/alkali pretreatment, sonication,
ionic liquid, etc.) also produces other lignin characteris-
tics (Table 1). Del Río et al. [17] analyzed this conception
with the comparison of milled wood lignin of angiosperms
with the milled gymnosperms. The comparison revealed
that acylated lignin units, including mono- and eudicot-
yledons, were observed in the angiosperms but lacking
in the gymnosperms. As β-O-4 linkages in lignin can be
cleaved without much energy, the highly acylated lignin
is more digestible than the non-acylated lignin. Further, a
significant aspect of lignin chemistry and its linkages with
the carbohydrate are represented in Table 1 [18–33].
27 Page 4 of 29 Advanced Composites and Hybrid Materials (2024) 7:27

Table 1  Compositional variability of lignin composition in wood of gymnosperms and angiosperms

Lignin% H unit % G unit % S unit % β-O-4% Phenyl cour- Resinol% 5–5′ Dibenzodiox- Β-1′% References
marins % ocins%

Gymnosperms
Cycas revoluta 30–40 0.7 98.7 - 59.5 24.9 5.4 5.9 [18]
Pinus pinaster 23–30 20 80 - - - [19, 20]
Pinus taeda 28 [21]
Picea abies 27 69 18 10 3 [22]
Pinus sylvestris bark 27.9% [23]
Quercus suber 24 1 44 55 76 9 8 [24]
Cunninghamia lanceolata 32.36–35.28 - - - - - - - - [25]
Angiosperm
Eucalyptus globulus 18 80 2 14 0 2 [22]
Agave sisalana 5–15 89 2 4 0 5 [26]
Hibiscus cannsbinus 5–15 84 2 8 0 6 [26]
Musa textilis 5–15 94 0 0 0 6 [26]
Ananas lucidus 5–15 94 2 0 0 4 [26]
Corchorus capsularis 13.3 72 4 16 0 4 [27]
Triticum aestivum 15 6 64 30 75 11 4 3 2 [27]
Zea mays 15 17 34 49 - - - - - [27]
Quercus suber - 8 84 8 68 20 4 5 - [24]
Betula pendula 22–25 - - - - - - - - [28]
Acacia mangium 28 1 1 0.1 40 43 - - - [29]
Quercus suber 24 1 44 55 77 9 8 - - [24]
Cynara cardunculus 16–19 0 58 42 70 14 7 - - [30]
Pyrus communis 0 68.3 31.7 [31]
Betula 20–35 0.1 17.3 82.6 38.4 10.1 1.6 1.8 [32]
Bambusa vulgaris 23.8 7.0 53.1 39.9 38.0 4.0 4.0 - - [33]

3 Lignolysis of biomass by microbes
The initial step of lignin breakdown, termed “lignolysis,”
is conducted by microbes such as fungi and bacteria. While
fungus-lignin degraders are restricted to white-rot and
brown-rot fungi, bacterial lignin-degrading species from
various environments, including genera such as Streptomy-
ces, Pseudomonas, Rhodococcus, Amycolatopsis, Oceani-
monas, Comamonas, Pandoraea, and Paenibacillus [34–48],
have been reported recently (Table 2).
Interesting observations on the comparative study for
lignin degradation of both the species revealed that while
fungi are efficient in higher and broader lignin degrada-
tion (due to the more comprehensive range of ligninolytic
enzymes), bacteria are more specific to the type of lignin
[38]. In the former, lignin depolymerization occurs in two
processes, selective and non-selective. While the lignin and
hemicellulose degradation occurs in the particular pathway,
the non-selective pathway is predisposed towards the degra-
dation of both polysaccharides and lignin [49]. The system-
atic action of fungus in lignin degradation is known to be
its secondary metabolic process as it is not essential for its
survival, which is contrary to bacterial mechanisms wherein
lignin degradation is known as one of the primary metabolic
processes [50, 51]. Consequently, bacteria are believed to
be more robust lignin degraders in terms of their expression
and activity [51]. Nevertheless, the general mechanism of
lignin degradation by both fungi and bacteria banks on the
concoction of peroxidases, laccases, and ­H2O2-generating
oxidases [49]. Peroxidases are featured enzymes that act on
lignin and lignin fragments and are remarkable in their abil-
ity to attack lignin from a distance [46]. Peroxidases produce
radical mediators that can enter lignin’s refractory structure
and cause depolymerization [46]. According to the type of
catalytic reaction they catalyze, these enzymes are catego-
rized as dye-decolorizing peroxidases (DyPs), lignin per-
oxidases, manganese peroxidases, and versatile peroxidases
(VPs) (Fig. 2) [52].
Manganese peroxidases (MnPs) are heme-containing
enzymes that use catalytic cycles via reactive M ­ n3+ ions
to attack lignin edifices [46, 53]. MnP shows a vital role
in the initial lignin depolymerization process and, without
any mediators, mainly subsidizes the cleavage of only 10%
lignin having ether linkage at C-4 of guaiacyl group and
hydroxylation of C-2 and produces 2,4-hexadiene-1,6-dioic
acid [54]. Moreover, the MnP can oxidize the non-phenolic
substrate of lignin structure by using phenolic mediators,
i.e., veratryl alcohol and fatty acids [55]. Although fungi
Advanced Composites and Hybrid Materials (2024) 7:27 Page 5 of 29 27

Table 2  Production condition and activity of different lignin-degrading enzymes

Microorganisms Substrates Enzymes Optimal condition Enzyme activity References

Phanerochaete chrys- Corncob LiP Peptone (0.3%), yeast 16.8 U/gds [39]
osporium extract (0.2%), and
Tween-80 (0.1%)
Acinetobacter sp. n-Propanol LiP pH—2.0 ­ g−1
1.571 U m [40]
L-DOPA, guaiacol, Temp—60 °C
hydroxyquinone,
catechol, and veratryl
alcohol
Trichoderma velutinum H2O2 LiP pH—5.0 7.6 U g­ −1 [41]
ACR-P1 Temp—30 °C
Pseudomonas putida Benzoic acid DyP-type peroxidase pH—7.0 ND [42]
type B1
Serratia marcescens Benzoic acid Protocatechuate 3,4-diox- pH—7.0 ND [42]
ygenase catechol-
1,2-oxygenase
Serratia liquefaciencens Benzoic acid Protocatechuate 3,4-diox- pH—7.0 ND [42]
ygenase catechol-
1,2-oxygenase
Trabulsiella sp. Guaiagylglycerol beta- LiP, β-aryl ether cleavage pH—7.0 ND [43]
guaiacyl ether (GGE) enzyme Temp—30 °C
Klebsiella pneumonia, Phenol red, guaiacol MnP pH—8.1 2.8 U ­mL−1 ­min−1 [44]
Salmonella entrica Temp—37 °C
Laccase pH—8.1 1.91 U ­mL−1 ­min−1
Temp—37 °C
Acinetobacter sp. n-Propanol LiP pH—2.0 ­ g−1
1.571 U m [40]
Temp—60 °C
Klebsiella pneumonia, Phenol red MnP, laccase pH—8.1 ND [44]
Salmonella enterica, Temp—37 °C
Enterobacter aerogenes,
Enterobacter cloaceae
Auricularia auricular- Veratryl alcohol DyP pH—1.4 ND [45]
judae Temp-NM
Steptomyces viridosporus NM DyP NM ND [46]
Steptomyces coelicolor ABTS, DMP Laccase pH—4.0 350 mg ­L−1 [47]
Temp—30 °C
Thermus thermophilus Guaiacol and 2,2-azino- Laccase ND ND [48]
bis(3-ethylbenzothiazo-
line-6-sulfonate)

have been widely observed to secrete MnPs, the production
of this enzyme from the bacterial community such as Kleb-
siella pneumonia, Salmonella entrica, Enterobacter aero-
genes, Enterobacter cloaceae, and Bacillus sp. has also been
proven (Table 2) [39–48].
Lignin peroxidases (LiPs) are triggered by the presence
of ­H2O2 and attack lignin via intermediate radicals, wherein
it oxidizes lignin compounds by single electron separation
from ­H2O2 and form radical cations. Non-requirement of
mediators during oxidation reaction marks LiPs as a more
potent oxidizing agent than that of manganese peroxidase
and laccase [56]. VPs, in some way, combine the catalytic
strategy of MnPs and LiPs [53] and are primarily active
at acidic pH (optimal pH around 5 is required in M ­ n2+
oxidation while pH 3 is the optimum for aromatic compound
oxidation) [52]. DyPs are structurally and phylogenetically
different from other peroxidases [53] and occur in fungi and
bacteria [46]. Similar to VPs, they are mainly active at acidic
pH values [46]. DyP has been classified into four subfami-
lies such as DyPA, DyPB, DyPC, and DyPD. DyPA contains
a Tat-dependent signal and is employed on the periphery of
the cell, while DyPB and DyPC are intra-cellular enzymes
that depolymerise anthraquinone and other aromatic com-
pounds [57]. Recently, Vignali et al. [58] overexpressed
DyPB of Rhodococcus jostii in Escherichia coli and revealed
that it efficiently oxidized aromatic lignin monomers and,
additionally, cleaved guaiacyl glycerol-beta-guaiacyl ether,
a model molecule for β-O-4 linkages. Type D participates in
27 Page 6 of 29
Fig. 2  Three-dimensional structures of different lignin-degrading enzymes important for their catalytic activities
the decolorization of dyes and is present in fungal variants
of Basidiomycota and Ascomycota. Though confirmation on
DyPs shows they participate in the degradation of broad
lignin substrates and defense against oxidative stress, their
physiological role is still imprecise (Table 2) [59].
Besides peroxidases, laccases considered an “ideal
green” catalyst play a key role in lignin degradation. They
only break down phenolic lignin units because they have a
lower redox potential than peroxidases [34, 52]. However,
this property is reversed in the presence of redox mediators
Advanced Composites and Hybrid Materials (2024) 7:27
such as vanillin and p-coumarate present [53]. The laccase
forms a stable intermediate with the substrate in the pres-
ence of mediators like 2,2′-azino bis C3-ethylbenzothiazo-
line-6-sulphonic acid (ABTS) and 1-hydroxybenzotriazole
(HBT) and with the help of these mediators depolymerize
almost 80–90% of lignin structure [60, 61].
Moreover, ­H2O2-generating enzymes such as aryl alco-
hol dehydrogenase, aryl alcohol oxidase, glyoxal oxidase,
and quinone reductase are essential contributors to lignin
degradation. Aryl alcohol oxidase (AAO) mostly found
Advanced Composites and Hybrid Materials (2024) 7:27
in Agaricales, Aspergillus, Fusarium, and even in some
bacteria contributes the hydrogen peroxide required by
ligninolytic peroxidase enzymes for the depolymerization
of lignin. Throughout the driving reaction of AAO and
aryl alcohol dehydrogenase, hydrogen peroxide is formed
which is used in the ligninolysis process. Besides, it is also
reported to ease the generation of guaiacol and sinapic acid
by laccase during the ligninolysis process [62, 63]. Glyoxal
oxidase (GLOX) by oxidizing a variety of simple bicarbo-
nyl and hydroxy carbonyl compounds such as glyoxal and
methylglyoxal generates extracellular hydrogen peroxide
[64]. The genes responsible for coding GLOX enzymes are
mainly found in wood-degrading fungi such as plant patho-
genic symbiotic fungi and white-rot fungi [65]. Further,
quinone reductase (QR), which also acts as a mediator in
lignin degradation, has been recently identified to impart
a more significant role in supplementing LiPs. Repolym-
erization of lignin by random coupling of phenoxy radicals
and quinoid intermediates, a common phenomenon during
lignin degradation by LiPs, is observed to be limited by the
addition of QR [66].
Fig. 3  Sustainable applications
of lignin in different industrial
and biomedical sectors
Page 7 of 29 27
4 Lignin as a platform for different chemicals
Recently, the employment of lignin feedstock into fossil-
based chemicals for industrial and transport-based products
has received broad attention. The potential of lignin to be
converted into different industrial value-added products is
given by its macromolecular structure and its reliability as
a strong biobase for industrial products (Fig. 3). Numerous
review articles have highlighted the valorization techniques
of lignin leading to different value-added chemicals and the
general category of the products in these studies has been
broadly listed under the three major categories, viz., power/
fuel, macromolecules, and aromatics (Table 3) [67–75].
However, the impediments in production of these value-
added products is the chemical structure of lignin that is
not ideal for bioconversion, taking into account its ineffi-
cient fractionation, the poor lignin processibility, and lower
yields of target products [76, 77]. Hence, the present review
focuses on the recent advancements in the lignin process-
ing aspects that can pave the platform for higher yields of
industrial products from lignin as biomass.
27 Page 8 of 29 Advanced Composites and Hybrid Materials (2024) 7:27

Table 3  Different value-added products from lignin

Feed stock Solvent Catalyst Condition Products References

Organosolv lignin Acetonitrile-THF Vanedium complex Temp—80 °C Vanillin—0.78% [69]

Ethylactetate-THF Time—24 h Syrinngic acid—0.67%
Syringaldehyde—0.59%
Organosolv lignin Ethylacetate Vanedium complex and Temp—100 °C Bio-oil—575 Da (MW) [70]
organometallic catalyst Time—18 h
Air—0.8 MPa
Wheat alkali lignin Water/methanol/1,4 CuO, ­Fe2(SO4)3 and Temp—150 °C Monophenolic com- [71]
dioxane/ethanol NaOH Time—1 h pounds—17.42% wt
H2O2
Kraft lignin Methanol water or CuSO4; ­FeCl3 Temp—170 °C Vanillin, methyl vanillate [72]
­H2SO4 CuCl2, ­CoCl2 Time—1 h
Air—0.5 MPa O ­ 2
Enzymatic hydrolyzed NaOH solution LaCO3 Temp- 120 °C Vanillin, p-hydroxybrnzy- [73]
lignin Time—3 h laldehyde
Air—0.5 MPa O ­ 2
Beech wood lignin NaOH solution La/SBH-15 Time—5–30 min Vanillin 9.94% syringalde- [74]
Microwave irradiation hyde—15.66%
200 W
Rice straw/husk/maple Temp—550 ̴ 900 °C Benzene [75]
Toluene
Xylene

For instance, Luterbacher and co-workers worked
on a pivotal work of aldehyde stabilization of lignin
and successfully improved the efficiency of lignin
depolymerization. Further, the result demonstrated good
yield for the mixture of guaiacyl (G) and syringyl (S) mono-
mers, such as alkyl-substituted G/S, hydroxypropyl-substi-
tuted G/S, and G/S propane diones, via hydrogenolysis and
oxidation approaches [78, 79]. In their pioneering work,
Rahimi et al. [80] demonstrated the depolymerization
of lignin into syringaldehyde and vanillin via successive oxi-
dation-formylation-hydrolysis processes. Wang and co-work-
ers [81] formulated the alkyl-substituted arenes and hydro-
carbons by direct hydrodeoxygenation of lignin. Recently,
Liao et al. [82] established an integrated biorefinery
process for converting lignin into high yields of phenol. The
complete transformation of Csp2-Csp3/Csp2-O of lignin into
C-H bonds without side hydrogenation is also significant for
obtaining value-added products from biomass [81].
Nevertheless, this has not been realized, so the area has
not been explored much. Even so, recently, Meng and co-
workers [83] developed a robust, high-performance proto-
col, wherein the transformation of the Csp2-Csp3 bonds on
the local structure of lignin molecule was conducted using
HY zeolite supported RuW alloy catalyst hydrogenolysis
for the production of benzene. The transformation resulted
in a scalable process for converting 50 g of lignin to 8.5
g of benzene. Liu et al. [84] also worked on the commer-
cial aspects of polyhydroxyalkanoate (PHA) production
from lignin, utilizing a “plug-in processes” approach. The
“plug-in processes” method combined with ammonium
flux pretreatment addresses the predicament of lignin- or
carbohydrate-first scenarios by solubilizing lignin via low-
ering molecular weight and increasing hydrophilic groups
strategy. Further, it also aids in the breakage of β-O-4 and
β-5 lignin linkages and enriched H- and G- type lignin, pro-
moting the consumption of lignin and PHA synthesis with
its selling price at as low as $6.18/kg.
Overall, the advancements in lignin processing aspects
have the potential to metamorphose lignocellulosic biorefin-
ery with better economics and sustainability. The biorefinery
designs for the production of value-added products should
be targeted to improve lignin conversion using the lignin
processing complexities as a platform. This approach will
be valuable for upgrading lignin at minimal costs and higher
product yields. Moreover, this could lead to profitable and
sustainable biorefinery prospects.
5 Lignin as composites
Composites are built from a polymer matrix and a reinforc-
ing material or designed synthetic or natural fiber. The basic
function of the matrix is to protect the fiber from exter-
nal damages caused by the environment and share the load
between the fibers. On the other, the fiber provides strength
and stiffness to the matrix, thereby enhancing its resistance
to cracks and fractures. Nowadays, biocomposites derived
from natural resource have garnered significant interest in
the automotive industry, manufacturing, aerospace, sports
equipment, construction, and marine applications due to
Advanced Composites and Hybrid Materials (2024) 7:27
their non-toxic and biodegradable nature [85]. These green
composites are mainly created from sustainable resources
derived from agriculture and forestry, encompassing crops,
and agricultural by-products. These residues primarily con-
sist of cellulose, hemicellulose, lignin, and other components
[86]. Among the natural polymers, lignin has functional
groups with various available sites that can be physically/
chemically modified and provides an excellent material to
serve as biocomposites. For instance, the phosphorylation,
hydroxymethylation, and oxypropylation reactions of lignin
derivatives provide outstanding biobased polyols for the syn-
thesis of polyesters and polyurethanes [87, 88].
An increase in the molecular weight and thermal stabil-
ity of the lignin favors its application in the epoxy resin
industry. Composites from lignin have shown promise as
adhesives, resins, varnishes, plasticizers, and flame retard-
ants [89]. Native lignin loses its reactivity when utilized
directly, lowering co-polymer composites’ mechani-
cal strength. As a result, it can be altered before being
Fig. 4  Application of lignin in making lignin-based biocomposites and hybrid materials
Page 9 of 29 27
combined with polymers to produce better results [89].
Lignin nanocomposites are employed for foodstuffs and
fiber modification for textiles to improve adhesion, hydro-
phobicity, antibacterial, and antioxidative properties of
the material. Other applications of lignin nanocomposites
include their employability as adsorbents in water purifi-
cation, emulsifiers for colloids, carriers for enzymes, and
controlled-release vectors for medications and insecticides
[6]. Lignin can be used to produce a variety of lignin com-
posites, such as lignin-polypropylene, lignin-epoxy resin,
lignin-polylactic acid, lignin-cellulose-based composite,
lignin-polyvinyl alcohol composites, lignin-starch com-
posites, lignin-chitosan composites, and lignin-based rub-
ber composite (Fig. 4).
Thus, owing to its ecological friendliness and enhanced
biocompatibility, lignin can be explored for producing high-
value composites [90, 91]. The following subsections elabo-
rate on the different lignin composites that can be high-value
products in the future.
27 Page 10 of 29
5.1 Lignin‑polypropylene composites
Polymer nanocomposites (PNCs) have perceived signifi-
cant growth due to their remarkable performance, stemming
from their distinctive physicochemical attributes. These
composites are lightweight and cost-effective and showed
excellent thermal stability, strong flexibility, and adjustable
mechanical, magnetic, rheological, and electrochemical
characteristics [92]. Among the polymer nanocomposites,
polypropylene (PP) is an important material that is required
by the industrial sector for manufacturing car batteries, sen-
sors, kinesiology, aerospace vechicle anticorrosion coating,
microwave absorbers, and, importantly, as packing materi-
als [93]. The current market cost of PP is around $1210/t,
which makes it an expensive raw material for the industrial
platform. Further, PP also has the disadvantage of being
unable to break down naturally with poor mechanical quali-
ties [94]. Thus, a partial replacement of PP with cheap raw
materials such as lignin will be ecologically and economi-
cally attractive. For instance, Alassod et al. [95] examined
various lignin loading levels in polypropylene and used the
thermally induced phase separation (TIPS) method (which
allows dissolving a polymer in a suitable high-temperature
solvent followed by cooling of the polymer solution) to
create inexpensive and environmentally friendly sorbents.
The authors concluded that the lignin range of 10–20% by
weight, along with PP, allows the formation of low-cost,
eco-friendly, and stable sorbents for oil clean-up applica-
tions in water bodies. In a similar study, a recent report by
Pregi et al. [96] emphasizes the use of 50–80 wt % of lignin
and lower amounts of flax seed ratios for improving poly-
propylene’s carbon footprint significantly. The authors also
concluded an interesting finding that both lignin and flax
seed acted on PP for additive properties individually but
not synergistically. Though solid lignin has been a preferred
choice for the production of lignin-based PP composites,
“liquid wood” or industrial waste streams with high wood
waste have also been experimented with for lignin/PP com-
posites [97]. The study concluded that liquid wood, along
with 20% hemp, was sufficient to form PP-lignin compos-
ites that were able to sustain three recycling cycles. The
PP-lignin biocomposite also exhibited favorable retention
of mechanical properties and thermal stability through recy-
cling. Yan et al. [98] conducted a recent study to investigate
the effectiveness of microwave irradiation as an environmen-
tally friendly technique for producing electrically conductive
polymer nanocomposites containing a minimal amount of
carbon nanotubes. This well-structured network of carbon
nanotubes (CNTs) led to increased electrical conductiv-
ity, with the electrical conductivity mechanism revealing
a three-dimensional variable range hopping (VRH) behav-
ior. The electrical conductivity of polymer nanocomposites
(PNCs) increased as the bandgap (Eg) decreased, indicating
Advanced Composites and Hybrid Materials (2024) 7:27
that PNCs with better conductivity possessed a lower Eg
[98]. Uday et al. [99] investigated the use of polypropylene
fiber to enhance the geotechnical characteristics of black
cotton soil. This approach offers the additional advantage
of reducing the need for traditional binders and decreas-
ing the expenses associated with industrial by-products.
Diesel fuels derived from waste tube-tire (WTT) and waste
polypropylene (WPP) were identified by employing stron-
tium carbonate (SrCO3) as a catalyst in a catalytic pyrolysis
cracking process. The diesel exhibited significantly lower
sulfur content compared to conventional fuel [100]. The
aforementioned studies provide necessary inputs for the use
of natural and lignin-based PP biocomposites [101]. How-
ever, some other reported literature emphasizes either the
modification of lignin or the addition of additional materials
to form better lignin-PP composites. The potential reason is
that lignin in its original structure does not show miscibility
with commercial polyolefins. More developed methods such
as TRIPS seem promising from economic and environmen-
tally friendly perspectives in such scenarios. In the future,
lignin-PP complexes can be relied on developing more such
methods. Also, wood-based industrial waste streams can be
targeted for the formation of these biocomposites, thus play-
ing a synergistic role for both eco-friendly industrial waste
disposal and also economical raw material accusations.
5.2 Lignin‑epoxy composite resin
Epoxy refers to a type of synthetic resin that is known
for its strong adhesive properties, excellent durability,
and versatility. Epoxy resins are created by mixing two
components: a resin and a hardener. When these two
components are combined, they undergo a chemical
reaction that results in a cured, rigid material with
exceptional bonding characteristics [102]. Epoxy resins find
applications in numerous fields, serving diverse purposes
including coatings, adhesives, modeling compounds,
impregnation materials, high-performance compos-
ites, insulating materials, shape-memory polymers, and
the encapsulation and packaging of electronic devices
[102, 103]. However, petroleum polymer–based epoxy
resin causes environmental pollution as it is toxic and
non-biodegrdable adding to the growing waste manage-
ment challenge. In response to these challenges, there
is an increasing focus on the advancement of biobased
epoxy resins and alternative, environmentally sustainable
materials that lessen dependence on petroleum-derived
resources and mitigate their adverse ecological effects.
Researchers have recently directed their efforts towards
lignocellulose fibers, including banana, sisal, coconut,
and oil palm fibers, for use as polymer reinforcements due
to its biodegradability, cost-effectiveness, low density,
and recyclability nature [101]. A significant category of
Advanced Composites and Hybrid Materials (2024) 7:27
thermosetting resins extensively employed in the coat-
ings and adhesives industries is estimated to have a global
production of approximately 2 million tons in 2010. Amelia
et al. [104] synthesized epoxy-based adaptable networks
through the combination of a diepoxide monomer and
an oligosiloxane prepolymer containing aminopropyl
groups, leading to irreversible cross-linking, as well as
silanolate end-groups that engage in dynamic bonding. This
epoxy composite, with its capacity to adjust thermal and
mechanical properties, renders it highly suitable for various
applications, including incorporation into laminates for
industries such as biomedicine, aerospace, or automotive.
Similarly, improved flame retardancy and dielectric
properties of epoxy resins were formed by incorporating
benzyl(4‑fluorophenyl)phenylphosphine oxide (BFPPO)
in its structure. These benefits support the potential of
BFPPO as a flame retardant in the manufacturing of
high-performance EP materials suitable for advanced
electrical applications [105]. However, researches have
been conducted on the creation of modified, partially or
completely biobased epoxy resins. In this context, lignin
has emerged as a notable component in this development
of biobased epoxy resins as it is biodegradable, cost-
effective, renewable, and sustainable; enhances ther-
mal and mechanical properties; reduces toxicity; reduces
carbon footprint; and valorizes waste products from
the pulp and paper industry or agricultural residues
[106]. Lignin offers the flexibility for the production of
a variety of materials, including hydrogels, composites,
and thermoplastics [107]. However, the most promising
application is the fabrication of thermosetting materials
such as lignin-epoxy composite resins. Epoxy resins are
the most popular thermoset material that is widely used by
industries for making sports equipment, aircraft parts, and
other composite materials [108, 109]. Epoxy resins cost
around USD 6341/t, making them an expensive polymeric
material for industrial use. Moreover, the epoxy resins are
non-porous and become slippery when wet (challenging
to handle), causing respiratory illness and contributing to
environmental pollution [110]. Many biobased substitutes
have recently been explored to eliminate the quantity of
bisphenol A content in epoxy resins, subsequently reducing
its cost. Thus, lignin biomass with functional groups such
as carboxylic acids, phenols, and aliphatic alcohols, along
with its rigid polyaromatic structure, allows the introduction
of functionalities suitable for subsequent material syntheses
of these novel thermoset polymers [108]. One such example
is utilizing industrial alkali lignin to prepare low-carbon
and environmentally friendly biobased epoxy thermosetting
materials. Recently, a small- and large-scale glycosidation
process was implemented to synthesize lignin-based epoxy
resins. Fifty to 60% of kraft lignin was used for epoxy resin
formation that lowered the chemical contents of epoxy
Page 11 of 29 27
resins and proved to improve the thermal stability of these
materials [108].
Gouveia et al. [111] used the epichlorohydrin method in
the preparation of lignin-based epoxy resins. The authors
concluded that the lignin modification increases its reactiv-
ity towards hardness in comparison to pristine epoxy and
might contribute to the reduction in the use of petroleum-
based resins. In a similar study, Li et al. [112] reported a
formaldehyde-free lignin-containing adhesive with broad
applications from furniture manufacture in the production
of engineered wood. This improved lignin-based wood
adhesive could be a potentially highly viable alternative
to widely used formaldehyde resins. Zhen et al. [113] per-
formed a novel strategy to synthesize phenolated lignin-
based incorporated novolac epoxy networks. This lignin-
based epoxy resin exhibited high flexural strength and high
storage modulus and significantly improved its thermal and
mechanical performance. The composite’s adhesive tensile
strength was noticeably affected by the lignin content, which
ranged from 10 to 20%. The shear strength of the glue only
increased after 4 h of heating at 75 °C [6]. A recent report
by Lu and Gu [114] emphasizes on combustion behavior of
epoxy resin by reducing smoke releasing and increasing char
formation by using alkali lignin. In their study, glyoxal as
a green chemical agent in place of formaldehyde was used
to improve the combustion behavior. The above-mentioned
studies showed lignin as an important biobased raw mate-
rial for biological epoxy resin preparation. However, lignin’s
extraction and purification processes make biobased epoxy
resin preparation more complex. There are many challenges
associated with lignin’s inherent variability and compatibil-
ity with epoxy chemistry. Hence, focus on preparation of
epoxy resin directly from raw biomass, or a simple LCB
pretreatment method will be useful to overcome these
limitations. The biobased epoxy resin still requires fossil
fuel–derived chemical bisphenol A. If epoxy resin prepara-
tion is entirely focused on renewable sources, it will promote
adequate biobased epoxy resin preparation.
5.3 Lignin‑based rubber composite
Rubber is a versatile material with a wide range of appli-
cations such as medical equipment, footware, tire manu-
facturing, medical equipments, electrical insulation, con-
sumer products, and other industrial applications [115].
Rubber composites are a special class of materials that
have helped industrial progress by creating several products
[116]. It provides unique behavior such as inherent capa-
bility, energy absorption, solvent resistance, high tensile
strength, and elastic recovery even at large deformations,
fillers, and other compounding ingredients. Yan et al. [117]
prepared boron nitride nanosheets and used a shear-coating
process to create composite films with enhanced thermal
27 Page 12 of 29
conductivity and electrical insulation properties. These
composite films were generated by using two components
of room-temperature vulcanized silicone rubber (RTV-2SR)
which were applied as a polymer matrix to enhance the ther-
mal conductivity of the rubber materials. This invention can
offer a fresh perspective on the development of heterostruc-
tured thermally conductive fillers and flexible composite
films that are thermally conductive and electrically insulat-
ing. However, the product formed is costly and uses petro-
leum-derived products for its preparation and formed materi-
als were non-environment-friendly. Similarly, carbon black
rubber material has been a known reinforcing filler in rubber
industries in the recent years [118]. In recent times, graphene
is being employed as a reinforcement in rubber composites
owing to its exceptional mechanical and thermal characteris-
tics. Graphene-rubber composites provide improved mechan-
ical and thermal characteristics; however, they encounter
obstacles such as elevated production expenses, challenges in
achieving uniform dispersion, specialized processing needs,
environmental considerations, and a lack of standardized pro-
cedures. Continuous research efforts aim to alleviate these
limitations and expand the utility of these materials [118,
119]. Thus, it is necessary to develop a non-petroleum-based
greener rubber composites due to environmental concerns.
Setua et al. [120] introduced lignin as a filler in nitrile rub-
ber composites. However, in the recent 5 years, lignin-based
rubber research has progressed speedily [118]. Lignin has
undergone silylation modification to improve its hydropho-
bicity and, consequently, its dispersion in the rubber system.
A 44.4% increase in tensile strength can be obtained by add-
ing 5% weight silylated KL to the natural rubber matrix.
The elastic moduli and Payne effect intensity increase with
the amount of modified lignin in the filler [121]. Kruželák
et al. [122] formulated rubber composites by incorporating
lignosulphonate, acrylonitrile-butadiene rubber (NBR), and
styrene-butadiene rubber. Glycerol was employed as a cost-
effective and eco-friendly plasticizer to enhance adhesion and
homogeneity between the rubber and filler materials.. From
the above studies, the kraft lignin-rubber composites with
higher thermal stability seem to be more effective compos-
ites compared to other rubber composites. Moreover, some
modifications would be useful to improve their stability in
making electrical and energy storage devices.
5.4 Lignin polysaccharide composites
Polysaccharides are macromolecules found in ani-
mals, plants, and microorganisms, forming structures
like glycogen and chitin in animals. They play a cru-
cial role in mineralization processes, providing a tem-
plate for mineral precursor orientation [123]. In recent
years, research in the fields of material science and
engineering has concentrated on polysaccharide-based
Advanced Composites and Hybrid Materials (2024) 7:27
composite materials because of their biocompat-
ibility, reusability, and sustainability. Numerous poly-
saccharides have garnered extensive research inter-
est because of their potential applications in various
fields, including tissue engineering, drug delivery, and
biopolymer-based film packaging [124]. Carrageenan,
chitosan, pectins, starch, and cellulose are main polysac-
charides that are used for making composites. Guo et al.
[124] created and characterized pectic polysaccharides
from hawthorn. It was done by using high methoxylated
pectic polysaccharide HSP which was acquired through
a gentle acid extraction process, utilizing hawthorn wine
residue as the primary raw material. In terms of function-
ality, HSP displayed a relatively elevated apparent viscos-
ity in an aqueous solution at various concentrations, yet it
did not exhibit gel formation. However, the polysaccha-
ride matrices are mechanically weak, with poor thermal
and vapor barrier properties. Various approaches were
made to enhance the properties of polysaccharide compos-
ites by adding inorganic plasticizers and organic fillers.
Organic and plasticizers generally use harmful chemicals
that are non-biodegradable and costly and cause pollution
to the environment. Thus, the replacement of this harmful
chemical with cost-effective, abundantly available lignin
will be more effective due to its high thermal stability and
biodegradable nature. Sadeghifar et al. [125] created and
characterized semi-transparent flexible cellulose-lignin
UV-light-blocking films. It was done by regenerating cel-
lulose-lignin films in acetone. These covalently bonded
cellulose-lignin films were homogeneous instead of sim-
ple mixes of cellulose and lignin. The efficacy of the
lignin polysaccharide complex as an antioxidant and UV
shielding material was investigated by Fu et al. in 2023.
The study concluded that lignin-polysaccharide compos-
ites exhibited a higher capacity of radical scavenging after
preactivation step of UV radiation and can be used to
develop oxidant and sunscreen products [126]. Wang et al.
[127] prepared lignin-polysaccharides using chitosan for
food packaging and tissue engineering materials. But
there are several drawbacks to this composite utilization
as it is costly, has low material resistance and poor sur-
face area, is acidic in nature, and has poor thermal stabil-
ity. However, adding lignin to the composite increased
solubility and improved mechanical strength [128]. The
composite hydrogel was prepared using lignin chitosan
for dressing the wound. Experimental work proves that
hydrogels’ mechanical strength increased when lignin was
introduced [129]. Çalgeris et al. [130] developed a biofilm
for drug release study by solvent casting process followed
by heating and drying using lignin, starch, and glycerol.
In their research, they found that acidic pH accelerated the
release of drug. These biofilms may also be used in many
areas, such as food packaging and drug delivery systems.
Advanced Composites and Hybrid Materials (2024) 7:27
6 Lignin hybrid materials
Recent research has shown a considerable increase in
interest in mixing lignin and organic polymers of natural
origin with silica to create novel hybrid materials. Future
synthetic polymers may be replaced by biodegradable
hybrid materials made from renewable resources, which
are incredibly promising sources. Inorganic (titanium,
zinc, aluminum, cobalt, iron, and silicon) and organic
hybrids such as lignin/chitin recently become intense sci-
entific research as they display several distinctive qualities
that have grown in significance in both the scientific and
technological fields [131]. Such hybrid materials exhibit
improved physicochemical, thermal, and mechanical
properties. Combining naturally occurring lignin with
silica, which has great mechanical and thermal resistance,
yields a hybrid that works well as filler in polymer sheets
to make a product with the needed qualities [132]. By
initially treating lignin with urea, other researchers
have tried to create a polylactide polymer with the best
possible flame-retardant qualities and mixing it with
ammonium phosphate [133, 134]. Culebras et al. [135]
investigated the fabrication of hybrid nanostructures
comprising lignin-derived carbon nanofibers (CNFs) and
silicon (Si) for potential use as high-performance anodes
in lithium-ion batteries. The utilization of a chitin/lignin
material and aminated lignin as an efficient adsorbent of
environmentally hazardous metals from model systems
was made possible by its well-defined physicochemi-
cal and morphological features. Its application in the
neutralization of genuine industrial wastes is particu-
larly significant [136]. The presence of multiple active
functional groups from the component biopolymers on
the surface of the newly produced product confirmed its
high sorption capacity when the ions ­Ni+2, ­Cu+2, ­Zn+2,
­As+5, and P ­ b+2 were adsorbed on the functional sorbent
[137]. The literature is replete with lignin being used to
stabilize polystyrene and polyethylene, with its primary
purpose being to guard against UV deterioration because
of its superior antioxidant capabilities. To avoid the
degradation of lignin during extrusion, future research
should concentrate on more lignin-based hybrid materials.
Numerous fields have shown significant development
potential from the above intensive research on lignin-
based composites and hybrid materials. More work needs
to be done on lignin-based polymer composites as these
are environment-friendly, cost-effective, available in huge
amounts, and biodegradable than petroleum base com-
posites. One of the major hindrances in the production of
lignin-based composites is its indistinctly defined struc-
ture, its quality differs along with its origin, and separation
and fragmentation using physical and chemical processes
Page 13 of 29 27
are costly. Biological methods use ligninolytic microor-
ganisms with other lignin-degrading communities, and
acquaintance with the production routes of these enzymes
can be beneficial in the fragmentation and separation of
lignin. The structural modification of lignin using bio-
logical methods partially or fully can enhance the reactiv-
ity, solubility, and compatibility of lignin in the produc-
tion of composite material. Production of uniform lignin
products via pathway may be useful in producing specific
lignin composites. In producing lignin-based composites,
various corrosive chemical activators cause corrosion in
the instruments. So, we use a green activator or design a
method without a corrosive activator to prepare lignin-
based composites.
7 Patent exploration for linin composite
and hybrid materials
Lignin-based materials hold a noteworthy place in the mar-
ket. The global lignin-based product market size is presently
valued at USD 1094.52 million in 2022 and is expected to
reach USD 1681.24 million by 2028 [138]. Patents form an
integral part of the market as they have a larger scope for
possible industrial applications. This section highlights the
published patents of lignin composites and hybrid materials
and its biomedical and non-biomedical applications during
the years 2010–2022 (Table 4).
The USA, China, and Europe are the top nations with
patents on lignin-based materials that will be helpful for
guiding future research. For example, the invention of bio-
degradable polymer formed by interlinking lignin with
butylene succinate leads to the production of biodegrad-
able polymer that reduces plastic pollution, carbon foot
print, dependency on fossil fuel, greenhouse gas emissions,
toxicity, and compostable [139]. Similarly, Tang et al.
[140] developed an innovative method for converting
thermoplastic phenolic resin into thermo-lignin phenolic
resins which offer many environmental and application
advantages over the former. The recent inventions on pat-
ents are focused on preparation of lignin compatible poly-
mer in which the lignin and polyol polymer are used [141].
Osterberg et al. [142] produced lignin-based hydrogels that
reduce the environmental impact by decreasing the rate
of chemicals used for making hydrogels apart from being
cost-effective. The comprehensive applications of the pat-
ents are presented in Table 4 [143–161]. The lignin-based
patented, specifically, biodegradable, compostable recy-
clable polymers are attractive products that can be indus-
trialized for replacing plastic-based materials. However, at
present, the processing of lignin-based polymers, though
Table 4  Patents related to innovative studies on lignin composites and hybrid materials
No Product name Manufacturing methods Application Patent type References

1 Lignin polymer nanocomposite Clay and lignin composites are melted and Showed good mechanical and thermal proper- Korean patent [160]
mixed with lactone compound ties KP20150061746A
27 Page 14 of 29

2 Nanocellulose-lignin composite Chemically associated with cellulose nanocrys- Cheaper material than petroleum composite World Intellectual Property and Organization [159]
tals and cellulose nanocrystals and physically having great mechanical and reinforcing WO/2015200584 A1
associated with lignin perporties
3 Lignin-based hydrogel Colloidal lignin particle dispersed in a mixture It showed shielding effect from ultraviolet light World Intellectual Property and Organization [142]
of organic solvent of lignin having excellent compatibility WO2020/109671A1
4 Carbon fiber composites Organosolv lignin used for carbon fiber com- High tensile strength and cost-effective renew- US patent [161]
posite able carbon fiber US011512173132
5 Lignin coating composites Lignin coating with a polygenic epoxy glycidyl Food and beverage packaging World Intellectual Property and Organization [150]
solvent WO2014/095800A1
6 5,4-Ethylene dioxy thiophene and lignosul- Lignosulfonic acid added to poly hydroxyl High conductive performance, high harness, China patent [144]
phonic acid containing composites aromatic compound high adhesion, heat stability, temperature CN102516784B
stability, and water resistance
7 Lignin thermoplastic phenolic resin Lignin phenolic compound, activated carbon, Improve reactivity of lignin and substitute rate China patent [140]
and aldehyde compounds are used for the of lignin on phenol CN102875752B
preparation of resin
8 Lignin-based hydrogel Lignin hydrophilic polymer and phenolic The water absorption rate increased Chinese patent [154]
hydroxyl group are interlinked with hydro- CN103834039A
philic polymer with epoxy group of ally
glycidyl ether and acrylamide
9 Lignin composite material Vinyl group, methacryloyl group grafting on The mechanical strength of composite improved World Intellectual Property and Organization [115]
lignin made to prepare composite WO2014/070036A1
10 Wood fiber–based plastic Lignocellulosic material mixed with DMSO The preparation method is environment- Chinese patent [157]
and plastic were prepared friendly, simple, and easy to operate CN103773054A
11 Lignin/polyacrylate compound oil absorbing Lignin acrylated A is prepared and acryloyl It is biodegradable material that can be used to Chinese patent [158]
material chloride is used to make this material remove oil from oil contaminated sewage CN103772627A
12 Lignin polyether polyols Alkali lignin, cellulose, sodium hydroxide, It is a novel foam material Chinese patent [155]
furfural alcohol, and furfural are used for the CN103772693A
preparation of this material
13 Lignin rubber composite Rubber modified using lignin Improve mechanical properties of rubber and Chinese patent [147]
reduce its cost CN103756060A
14 Lignin polyester polyol compound Calcium lignosulfate, water, cellulose, sodium Suitable starting material as making foam board Chinese patent [156]
hydroxide, furfural alcohol, and DEGC are CN103755974A
used
15 Lignin containing rubber composites Lignin and rubber are used Reinforcing fiber a green composite US patent [151]
US008664305B2
16 Lignin epoxy resin Lignin epoxy resin without any organic com- Used for coating, flooring, additives, lubricants, US patent [152]
pounds and thermosetting resins US20140039146A1
17 Lignin thermoplastic phenolic resin Phenol and enzymatically hydrolyzed lignin Green resin China patent [146]
used CN103554400A
Advanced Composites and Hybrid Materials (2024) 7:27
 Advanced Composites and Hybrid Materials (2024) 7:27 Page 15 of 29 27

possible, cannot be industrialized due to the complexity of the

References process. Hence, more research can be targeted towards these
[153] for production of economical and environment-friendly

[145]

[143]

[141]

[139]

[149]
lignin-based products.
World Intellectual Property and Organization

8 En route for lignin biotechnology

Commercialization of lignin for obtaining value-added prod-

ucts such as bio-oil (lubricant), carbon nanofiber, anode
material, and hybrid nanomaterials faces numerous chal-
WO2014/012924A1

elastomers, binders, thermoset resins, thermo- US20200140624A1

US20120071591A1
lenges [162]. The significant challenges are the necessity
US008747709B2
CN103524709A

European patent
EP2619251A1

of high quantities of lignin and the requirement being met

China patent
Patent type

US patent

US patent

US patent

mostly from contaminated lignin that is recovered from the

industries [163]. The contaminated lignin is mostly the sul-
fonated forms from kraft and sulfite lignin processes that
Hydrogenated enzymatic lignin was mixed with Biodegradable epoxy resin having low in pollu-

also comprises crude mixtures of various aromatics, some of

Used for making coating, adhesives, sealants,

It is a biodegradable, compostable recyclable

which may be toxic [164]. The breakdown of this contami-

Renewable source used for flame retardant
plastic resins, and polyurethane systems
Increase the mechanical reinforcement of

Precursor fiber having thermal resistant

nated lignin with the help of microbes is another challenge

from the biorefinery prospective. From the other prospec-
tive, microbial degradation of the tainted lignin entails alter-
ing their genetic components for enhanced stress tolerance
[165, 166]. Further, the broad spectrum of substrate, engi-
elastomer polymer

neering the routes of higher enzyme production for aromatic

tion and cost

catabolism, and rationalized pathway regulation are some

Application

polymer

material

of the other favorable characteristics in engineered micro-

organisms that have been investigated [167–169]. Never-
theless, metagenomics and synthetic biology engineering
approaches are attractive platforms for achieving the desired
Lignin having covalently linked with phospho-
Chemically unmodified lignin used in produc-

Thermoplastic lignins are used for producing

strain performance.
Lignin and butylene succinate are used
Lignin and polyol polymer are used

8.1 Metagenomic studies in lignin

tion of the novel biopolymer

Metagenomics, or environmental and community genom-

epoxy epichlorohydrin
Manufacturing methods

rus containing groups

ics, is a culture-independent approach where direct analysis

of environmental samples is performed that can be utilized
carbon fibers

to define a microbiome in two different levels: structural

and functional. It is an effective method for describing/
discovering the enzymes, metabolic pathways, and
microorganisms involved in lignin degradation. In addition
to their ecological interactions and evolutionary features, a
Phosphate-substituted lignin flame retardant

microbial community’s structural aspects are investigated,

from the phylum to species level [170, 171]. On the other
hand, functional metagenomics investigates the genetic
diversity of the sample’s bacterial population to identify
Lignin-based plastic material
Lignin compatible polymer

the genes and metabolic pathways common in a microbi-

Lignin-based elastomer

ome (Fig. 5). 16S rRNA gene amplicon and metagenomic

Lignin carbon fiber
Lignin epoxy resin

shotgun sequencing are the primary technological methods

Table 4  (continued)

for profiling microbial populations [172]. The use of

Product name

material

metagenomics, metatranscriptomics, and metaproteomics to

analyze environmental materials directly is a strong technique
for describing/discovering enzymes, metabolic pathways
(Fig. 5), and microbes involved in lignin degradation. In
No
18

19

20

21

22

23
27 Page 16 of 29
fact, applying these complementary approaches enhances
knowledge of the makeup, characteristics, and dynamics
of microbial communities engaged in lignin deconstruction
[173]. In their study, Moraes et al. [174] analyzed lignin-
degrading consortia (LigMet) isolated from a soil sam-
ple from a sugarcane plantation. Based on 16S rRNA, the
LigMet taxonomical analyses revealed that Proteobacteria,
Actinobacteria, and Firmicutes members including those
from the Alcaligenaceae and Micrococcaceae families
were more common and concentrated in the LigMet than in
sugarcane soil. Over 240,000 gene models were discovered
through analysis of global DNA sequencing, and 65 draft
bacterial genomes were anticipated.
The main pathways for aromatic degradation, such as
benzoate (or methyl benzoate) degradation to catechol (or
methyl catechol), catechol ortho-cleavage, catechol meta-
cleavage, and phthalate degradation, were identified along
with several peroxidases, dye-decolorizing peroxidases, lac-
cases, carbohydrate esterases, and lignocellulosic auxiliary
(redox) activities. LigMet identified a novel Paenarthro-
bacter strain that could thrive on lignin as its primary car-
bon source and had eight gene clusters relevant to aromatic
degradation. Additionally, using novel gene sequences for
an enoyl-CoA hydratase/aldolase and feruloyl-CoA syn-
thetases that were found in the metagenomic data collection,
a recombinant pathway for the manufacture of vanillin was
developed [175].
Scully et al. [10], in their study, revealed the metagenomic
profiling in a microbial community associated with a wood-
feeding beetle (Anoplophora glabripennis) responsible for
lignocellulose degradation. The A. glabripennis gut metage-
nome contained 36 families of glycoside hydrolases, includ-
ing cellulases and xylanases, candidate lignin-degrading
genes (laccases, dye-decolorizing peroxidases, novel peroxi-
dases, and -etherases), and genes that could promote nutri-
ent recovery, essential nutrient synthesis, and detoxification.
These genes could also play important roles in the digestion
of woody tissues. This community may act as a repository
for cutting-edge enzymes to improve the production of cel-
lulosic biofuels in the industrial setting or as potential targets
for cutting-edge insect control techniques. In their inves-
tigation, Levy-Booth et al. [175] performed the genomic
and metatranscriptomic analyses of biochemical cycling
and degradation of lignin-derived aromatic compounds in
thermal swamp sediment. The study revealed an active role
of temperature in maintaining biogeochemical cycling and
lignin degradation in microbial consortia. A total of 351
distinct genomes from hot spring (30–45 °C) and mesophilic
(17 °C) sediments were resolved after the assembly of shot-
gun metagenome libraries. Annotation of 39 genomes per-
formed in this work indicated oligotrophy-like metabolism,
including aromatic compound breakdown pathways such as
syringate, vanillate, p-hydroxybenzoate, and phenol. Brink
Advanced Composites and Hybrid Materials (2024) 7:27
et al. [176], in an attempt, tried to bridge the gap between a
basic understanding of microbial lignin catabolism and the
newly developing field of lignin biovalorization via applied
biotechnology. The eLignin Microbial Database (www.e​ lign​
indat​abase.​com), an open-access database that indexes data
from the lignin bibliome, including bacteria, aromatic sub-
strates, and metabolic pathways, has been created for this
purpose. Wilhelm et al. [177], in their study, performed the
metagenomic analysis and quantitative stable isotope prob-
ing (SIP) for the establishment of bacterial contributions to
lignocellulose degradation in forest soils. In this study, SIP
significantly improved the shotgun metagenome assembly
by which several high-quality draft metagenome-assembled
genomes were recovered, as well as approximately 7500
contigs, including unique clusters of carbohydrate-active
genes, were identified. Using 454 pyrosequencing of 16S
rRNA, Su et al. [178] examined the microbiome gut of four
species of termites (three species of wood-feeding termites
and one species of fungus-feeding termites). Spirochaetes
(11–55%), Firmicutes (7–18%), Bacteroidetes (7–31%),
and Proteobacteria (8–14%) were found to be the primary
phyla for all four termites in this analysis. These findings
demonstrate the presence of genes encoding lignin modi-
fication or breakdown enzymes in termite gut microorgan-
isms, despite the fact that their function in the termite gut is
unknown [179]. Ufarté et al. [180] discovered that phenolic
compounds like ABTS (2,20-azino-bis(3-ethylbenzothia-
zoline-6-sulfonic acid)), syringaldehyde, 1-hydroxybenzo-
triazole, acetosyringone, and 2,2′-azinobis can be oxidized
by enzymes in the bovine rumen microbiome without the
addition of mediators like copper or manganes (4.5–5.5).
Using a bovine rumen metagenomic library, Beloqui et al.
[181] revealed RL5, a gene encoding a polyphenol oxidase
with laccase activity. This laccase sequence was assigned to
the genus Bacteroides by phylogenetic analysis. Agamen-
none et al. [182] used shotgun sequencing and bioinformat-
ics methods to examine the gut microbiome connected to
the soil invertebrate Folsomia candida. In total, 2004 genes
encoding enzymes that break down cellulose, starch, and
lignin were discovered. From these, 1988 (99.2%) were
primarily protists (43%) and actinobacteria (36%), which
together accounted for 99.2% of the bacteria. There were 81
genes identified within the CAZyme families as auxiliary
activities, with the AA3 family members accounting for 32%
of the total, AA6 for 27%, and AA7 for 17% of the total. The
range of ancillary activities discovered indicated that the
bacterial microbiome might contribute to the invertebrate’s
adaptation to soil life by degrading or altering lignin.
Metagenomic is a pioneering research based on numer-
ous unrevealed novel lignin-converting enzymes, aromatic-
degrading bacterial genomes, and a unique bacterial strain
with biotechnological promise. Targeted metagenome dis-
covery technique for lignin valorization strategies has been
Advanced Composites and Hybrid Materials (2024) 7:27
Fig. 5  Metabolic pathway of lignin degradation
used extensively to study other biosynthetic pathways that
help produce various lignin-derived products. The focused
metagenomic discovery platform can potentially uncover
optimal gene sets and microbes for synthetic biology-based
projects to manufacture high-value compounds from ligno-
cellulose. Understanding from metagenomic can increase
knowledge of which species, circumstances, and functional
genes are involved in lignocellulose breakdown. Apart
from this, the metagenomic method might be used in vari-
ous domains, including the creation of antibiotic-producing
bacteria and the recycling of plastic polymers.
8.2 Role of synthetic biology in engineering
microbes for enhanced lignin utilization
Lignin-degrading microorganisms use different fun-
neling pathways to channel different lignomonomers
through the essential intermediates. The non-conventional
Page 17 of 29 27
microorganisms that exhibit these properties are challeng-
ing to engineer. The synthetic biology tools in this scenario
are thought to play a major role in the hosts, such as bac-
teria, fungi, and unicellular yeast for the biotransformation
of lignin. Many reviews have represented excellent data
on various catalytic manipulation methods for converting
lignin [10, 14, 17, 183]. Hence, the proceeding sections
will discuss the recent advancements in the system biology
approach considering the microorganisms mentioned above
as hosts.
8.2.1 Bacteria as a host
Bacteria are more acquiescent to genetic engineering than
fungal and yeast cells [184]. Ece and co-workers [185]
cleverly expressed codon-optimized genes from Escheri-
chia coli under a cold-induced promoter (cspA) along with
the co-expression of chaperone DsbC in Ceriporiopsis
27 Page 18 of 29
subvermispora. A soluble, active manganese peroxidase
(MnP) with a properly incorporated heme resulted in a 30
mg/mL protein titer. E. coli chaperones coupled with ATP
and hemin were expressed in Phanerochaete chyrosporium,
producing MnP comparable to the commercial titers [186].
Other groups have supplemented more efficient secretion
tags to escalate bacterial laccase expression in non-E. coli
bacterial systems. For instance, heterologous expression of
a laccase that was adopted from Streptomyces coelicolor
was enabled in Rhodococcus opacus PD630. The system
was engineered at its promoter, ribosome binding site
(RBS), and an upstream region of laccase, i.e., secretion tag,
along with a Tat secretory system. The engineered PD360
showed enhanced production of laccase and it facilitated
the degradation of 81.1% of β5 linkages, 40.8% of aliphatic
hydroxyl group linkages, and 26.3% of the guaicyl hydroxyl
group as compared to the laccase negated wild strain lead-
ing to a total degradation of 33.9%, 15.5%, and 8.9% of
the aforementioned linkages respectively [187]. In a simi-
lar work, overexpression of DyP from Amycolatopsis sp.
was conducted on Pseudomonas putida using an effective
promoter ­(P1099) and an endogeneous secretion tag (PelB).
Engineered P. putida exhibited an enhanced growth (2.1-
fold) in kraft lignin and more than 40% reduction in the
β5 phenolic groups compared to the native strains. Even
though bacterial platforms have the advantages of shorter
doubling times and wide adaptability, the expression of fun-
gal enzymes by bacteria possesses some drawbacks. The
limitations of insufficient secretion mechanisms, the non-
existence of post-translational modification, and the skiving
of chaperones and cofactors lead to unsuccessful folding
and lower protein performance. On the other hand, although
yeasts have some other limitations, they are apprehended
to be a better platform for the synthetic engineering of lac-
cases and peroxidases.
8.2.2 Yeast as a host
Detoxification of pulping black liquor by expression of
POXA 1B laccase from Pleurotus ostreatus by using Pichia
pastoris as a host was conducted by Rivera-Hoyos and col-
leagues [188]. The essential features such as the chemical
oxygen demand (COD) and the decolorization of synthetic
black liquor (SBL) were evaluated for both the engineered
and natively expressed enzymes coupled with CuO/TiO2/
visible-light photocatalysis. The recombinant P. pastoris
demonstrated 63.7% COD removal along with 46% decolori-
zation of SBL, which was impressive, but not at par with the
performance of wild fungal P. ostreatus that showed 80.3%
and 70.6% COD removal and decolorization respectively.
The decreased catalytic activity may be due to the hypergly-
cosylation reaction in P. pastoris. Mateljak et al. [189] used
Saccharomyces cerevisiae as a host for expressing OB1 from
Advanced Composites and Hybrid Materials (2024) 7:27
Coriolopsis sp. PM1, 3PO from Pyconoporus cinnabarinus,
and Lac3 from Trametes sp. strain C30 through a structure-
guided SCHEMA approach for producing recombinant vari-
ants. The variants showed higher catalytic efficiency and
enhanced pH stability in acidic conditions alongside higher
thermostability. Song and his co-workers [190] integrated
cDNA from P. ostreatus in P. pastoris. Recombinant P. pas-
toris demonstrated 18.36% degradation of corn stover com-
pared to 14.05% by native strain in the same experimental
conditions. Following the production of synthetic recom-
binants, further steps can be targeted for enhanced ligninase
production utilizing a multicopy plasmid approach (MPA)
[191]. The MPA aims for a dramatic rise in the copies of
transcription units that carry the enzyme coding sequence
(CDS) into the yeast cells [192]. Further, the host cells allow
the replication of episomal shuttle (as plasmids can replicate
both in bacteria and yeast) vectors (ESV) autonomously to
express heterologous genes in yeast [193]. Once the gene of
interest is inserted in the ESV in in vitro conditions, the bac-
terial and yeast cells are transformed with the newly assem-
bled plasmid. The symbiotic association targets bacteria as
a means of storage while yeasts arrange the mechanism for
gene expression.
8.2.3 Fungus as a host
Even though yeasts exhibit significant advantages over fungi
and bacteria, the challenge in the recombinant expression
of the ligninases fuels the urge to engineer fungal systems
for its enhanced production. Mekmouche et al. [193] used
Aspergillus niger to express lac3 from Trametes sp. C30 to
increase its catalytic activity for ABTS (2,2′-azino-bis(3-
ethylbenzothiazoline-6-sulfonic acid)), syringaldazine,
and guaiacol. Disappointingly, except for the increase in
its titer by 200-fold, the catalytic activity of the engineered
A. niger towards the aforementioned lignols decreased by
71%, 57%, and 54%, respectively, compared to the cata-
lytic activity of S. cerevisiae that was transformed with the
same gene. Li and his co-workers [13] incorporated pslcc,
a laccase gene from Pycnoporus sanguineus, into protease-
(DppV and PepA) deficient Aspergillus nidulans using an
activated promoter (­ PalfR). The engineered A. nidulans had
a 15-fold and 13-fold increase in its expression and total
enzyme activity respectively compared to its native strain.
The non-targeted lac gene integration strategy from Pyc-
noporus sanguineus was targeted for Trichoderma ressei in
a fed-batch operation that resulted in a final protein titer
of 17.7 U/mL and an increased thermostability. A similar
strategy of non-targeted multiple genes, i.e., MnP1, LiPH8,
Vpl2, and Laclllb from endogenous strain, Pleurotus erygnii,
and Trametes versicolor, respectively, was integrated onto
Phanerochaete chrysosporium. A notable increase in lignin
Advanced Composites and Hybrid Materials (2024) 7:27
depolymerization (25%) along with 2.6- and fourfold higher
peroxidase and laccase activity, respectively, was observed
compared to the wild P. chrysosporium.
The synthetic biology approach for engineering bacteria,
yeast, and fungi to produce value-added products from lignin
or enhance lignin depolymerization specifically by selecting
optimal hosts is an optimistic headway. However, the lack of
predictive understanding of the post-translational modifica-
tions that strongly influence the enzyme activity makes the
progress of synthetic biology approaches dawdling. Addi-
tionally, very little work has been done on the synthetic
biology tool set for fungal systems with significantly higher
catalytic activity than the heterologous bacterial and yeast
systems. Hence, a thoughtful, intensive focus on the aspects
mentioned above will be helpful for the development of
more robust synthetic biology systems.
8.3 In silico study on ligninolytic enzymes
The enzymes are proteins that act as biological catalysts
with conserved functional groups. These conserved func-
tional groups play crucial functions in enabling the enzyme
to bind to its substrate and promote the catalytic reaction.
Several techniques like electrophoresis, NMR, X-ray dif-
fraction, and chromatography have been employed for pro-
tein purification and the enzyme’s physicochemical and
structural nature. But all these procedures consume more
time and money. Therefore, now, in silico approaches have
been applied to better understand the structure and func-
tional characteristics of an enzyme (proteins) substrate
interaction pattern in a cheaper, faster, better, safer, and
more scalable way. In silico study of enzymes deals with
the three-dimensional structure prediction, its validation,
and the survey of catalytic interaction purely using a com-
putational database [194]. The in silico study about ligni-
nolytic enzymes helps us to understand the 3D structures
of these enzymes, their interaction with the substrate, and
their validation on the study of catalytic activity using com-
puter stimulation approaches (Fig. 6). The ligninase protein
sequence found in the NCBI database revealed that lignin
peroxidase is a glycoprotein with a molecular mass of 38–46
kDa containing 1 mol of iron protoporphyrin IX per 1 mol
of protein in its structure. Another ligninolytic enzyme,
manganese peroxidase, is a glycosylated heme protein hav-
ing 40–50-kDa molecular mass. Laccase is a ligninolytic
glycosylated protein containing four copper ion binding
sites in its structure. It has different residual interactions
and the Cu binding motif shown to play a significant role
in the biosynthesis and degradation of lignin [195]. Singh
et al. [196] unraveled the lignin degradation mechanism by
performing in silico docking and simulations of lignin per-
oxidase enzymes with different lignin model compounds.
In this study, 30-ns molecular dynamic simulations were
Page 19 of 29 27
performed for the docked complexes of LiP enzyme with
different lignin model compounds, after which Gibbs free
energy of the complexes was calculated from the MD simu-
lation trajectories. The results from this investigation show
that bacterial LiP can efficiently catalyze a variety of lignin
model compounds and may be utilized as a tool for the
long-term mitigation of various environmental pollutants.
Díaz et al. [197] made an in silico design of a fungal
laccase, Lacc 6 from Pleurotus ostreatus, to make it more
thermostable. The acquired results from this study allow for
a better understanding of the molecular basis for increasing
the structural stability of the P. ostreatus Lacc 6, as well
as the in silico creation of mutants with high-temperature
activity. Mathé et al. [198] defined a new class of ligninases
by performing a high-quality gene annotation. In this study,
well-conserved gene clusters have been identified by phy-
logenetic analysis and compared with existing profiles to
construct new profiles and subprofiles. Pham et al. [199]
performed an in silico design of lignin peroxidase from
Phanerochaete chrysosporium, which exhibits enhanced
acid stability during the depolymerization of lignin. This
study suggested an in silico–based strategy to design active
LiPH8 variants for increased stability in an intensively
acidic environment. The structure of the LiPH8 variant was
solved by the molecular auxiliary method using MOLREP
based on the original LiPH8 (PDB Id-IB80) [200]. WinCoot
and REFMAC programs were used for model building and
refinement of the model structure [201, 202]. After execut-
ing the in silico mutagenesis, the salt bridge and MD simu-
lation studies showed more excellent stability under highly
acidic conditions, with a 12.5-fold increase in half-life and
a 9.9-fold increase in catalytic effectiveness towards veratryl
alcohol. Gibbs free energy of the LiPH8 variant protein was
also calculated computationally after introducing a differ-
ent mutation in the protein structure. The instability of the
intra-protein salt bridges was measured by calculating the
Gibbs, as mentioned earlier, free energy distribution [203].
In silico analysis of lignin peroxidase of the fungus Mac-
rophomina phaseolina (M. phaseolina) was studied by Alfi
et al. [186]. The 2D structure of lignin peroxidase was ana-
lyzed using SOPMA (self-optimized prediction method with
alignment tool). The result of 2D structure analysis showed
the predominance of the random coil followed by alpha helix
extended strand and beta-sheet, respectively [204]. The 3D
structure of lignin peroxidase was predicated using 3D-JIG-
SAW comparative modelling server [205]. Motif detection
and their location in the sequence were studied using motif
scan tool [206]. The motif detection result showed that lignin
peroxidase found in M. phaseolina and many other fungi
could degrade lignin efficiently [207]. Weirick et al. [208]
studied laccase enzyme class using in silico approaches.
This study offers a useful classification system of laccase for
understanding their physicochemical properties. This study
27 Page 20 of 29 Advanced Composites and Hybrid Materials (2024) 7:27
 Advanced Composites and Hybrid Materials (2024) 7:27
◂Fig. 6  In silico, metagenomic profile and synthetic biology study of
lignolytic enzymes for better understanding their interaction with sub-
strate and for searching targeted microbes having less defective enzyme
investigated several self-organizing map (SOM) architec-
tures to understand the effect of clustering of laccase with
many descriptors. Analysis of the taxa in each class revealed
that single taxa dominated most categories. Martinez et al.
[209] studied mapping the long-range electron transfer
(LRET) route in ligninolytic peroxidase enzymes using dif-
ferent in silico methods. In this study, ligand diffusion and
docking stimulation were performed with PELR (protein-
energy landscape exploration) for modelling ligands. The
LRET pathway in lignin peroxidase was also analyzed,
which showed that lignin peroxidase could oxidize differ-
ent aromatic compounds.
From the availability of comparative data for different
ligninase enzymes, it can be noted that the laccase enzyme
has the highest number of amino acid sequences and 3D
structure information. It can also be noted that bacteria
have a significantly greater number of laccase amino acid
sequences from various organisms. As with the laccase
enzyme, more lignin peroxidase amino acid sequences have
been found for bacteria. For manganese peroxidase, most
of the amino acid sequence for this enzyme is shown to be
retrieved from fungi.
9 Conclusion and prospective
In spite of substantial attention for conversion of lignin into
value-added products, less than 2% of lignin is utilized in
industrial scale, majorly due to constraints in the available
conversion technology. Considering the fact that the future
production of biofuels will be progressively inclined towards
lignocellulosic biomass, it is noteworthy that considerable
amount of residual lignin would be generated. The primary
step would be a thorough understanding of the structure and
component ratios of the derived lignin for their skillful utili-
zation. The present review briefs a considerate insight into
the different structure and components of different wood-
based lignin that will be helpful for the future lignin chem-
ists. Following that, the conversion technologies that form a
crucial part of the valorization of the lignin into value-added
products are also reviewed. The present conversion technolo-
gies that are based on chemical and physical methods might
not be attractive due to (a) non-specificity that might affect
the degradation of cellulose and hemicellulose, (b) economic
factors, and (b) environmental issue due to the by-products
generated by these products.
Biological conversion technologies involving enzymatic
degradation of lignin that has the advantage of specificity
and less-time consuming as compared to the other processes
Page 21 of 29 27
look promising. At present, in vitro enzyme-based degra-
dation of lignin compounds has been conducted using
well-known enzymes such as laccase, dye-decolorizing
peroxidase, manganese peroxidase, versatile peroxidase,
and lignin peroxidase. But these enzymes, moreover, have
been useful to break the specific linkage of lignin model
compounds and mostly lack the ability to degrade real
complex lignin residues. Thus, development of enzymes
mostly targeting the β-O-4-aryl ether linkage will be a
potential approach for adopting enzyme-based lignin
conversion methods. Adding to this, a better understanding
of ligninolytic microorganisms for production of higher titer
of lignin-degrading enzymes will form a crucial platform for
transformation of lignin into high-value-added products such
as chemicals and biocomposites.
The macromolecular structure of lignin allows it and
its derivatives to be a valuable platform for production of
many value-added chemicals such as vanillin, benzene, and
hydrocarbons to name a few. Apart from producing these
chemicals, lignin, that has a high carbon content, can be
considered an ideal carbon material precursor and used in
fields like energy storage, adsorbent, and catalyst carriers
[210]. Further, conversion of lignin and its derivatives into
biocomposites is a recent field of interest for many research-
ers. This review highlights the prospects of using lignin and
its modification strategies to serve as biocomposites. It is so
far inferred that native lignin loses its reactivity when uti-
lized directly, lowering co-polymer composites’ mechanical
strength. Therefore, it can be altered before being combined
with polymers to produce polyesters and polyurethanes.
However, reports also emphasize that lignin in its original
structure does not show miscibility with commercial poly-
olefins [211]. Thus, future application of thermally induced
phase separation methods seems promising from economic
and environmentally friendly perspectives in such scenarios.
Also, wood-based industrial waste streams can be targeted
for the formation of these biocomposites, thus playing a syn-
ergistic role for both eco-friendly industrial waste disposal
and also economical raw material accusations. Further, the
most promising application of lignin-based composites in
the future is expected as the fabrication of thermosetting
materials such as lignin-epoxy composite resins for mak-
ing sports equipment, aircraft parts, and other composite
materials. The present market for lignin-epoxy composite
resins faces the challenges associated with lignin’s inher-
ent variability and compatibility with epoxy chemistry.
Developments in procedures for preparation of epoxy resin
directly from raw biomass will be useful to overcome these
limitations. Also, at present, the biobased epoxy resin still
requires fossil fuel-derived chemical bisphenol A. If epoxy
resin preparation is entirely focused on renewable sources,
it will promote adequate biobased epoxy resin preparation.
Adding to it, increase in interest in mixing lignin and organic
27 Page 22 of 29
polymers of natural origin with silica to create novel hybrid
materials has also been observed recently. While research
on lignin-based hybrid materials are still on primary phase,
its application in the neutralization of industrial wastes is of
significant interest. For a practical observation of the use of
the lignin-based composites, this review also highlights the
significant patents that have been published over a decade.
Patents form a real-world approach towards the application
of the processes and the products on an industrial scale. It
is noteworthy that scalability of the lab-scale products is
rarely conceivable. However, patents do provide a biorefin-
ery based approach for development of products.
Development of lignin-based products in the future will
be strongly directed by genetic modification lignolytic
microorganisms. In this context, microbial engineering for
enhanced enzyme production for aromatic catabolism and
rationalized pathway regulation will be attractive platforms
for achieving the desired strain performance. The use of
metagenomics, metatranscriptomics, and metaproteomics to
analyze environmental materials for discovery of enzymes,
metabolic pathways, and microbes involved in lignin degra-
dation will hold a crucial role for lignin conversion strate-
gies. Application of these approaches will augment infor-
mation of the makeup, characteristics, and dynamics of
microbial communities engaged in lignin deconstruction.
For instance, Levy-Booth et al. [175] performed the genomic
and metatranscriptomic analyses of biochemical cycling and
degradation of lignin-derived aromatic compounds in ther-
mal swamp sediment and concluded that temperature has
an active role in maintaining biogeochemical cycling and
lignin degradation in microbial consortia. Metagenomic
discovery platform can potentially uncover optimal gene
sets and microbes for synthetic biology-based projects to
manufacture high-value compounds from lignocellulose.
Understanding from metagenomic can increase knowledge
of which species, circumstances, and functional genes are
involved in lignocellulose breakdown. Further, other biologi-
cal tools such as application of synthetic biology approach
for engineering bacteria, yeast, and fungi to produce value-
added products from lignin or enhance lignin depolym-
erization specifically by selecting optimal hosts is also an
area that can be explored in the future. While bacteria have
been the most preferred host till date because of the ease
to of genetic manipulation in it as a host, the low enzyme
productivity still remains a hurdle. Conversely, fungal/
yeast species, that naturally yield higher titers of lignolytic
enzymes as compared to bacteria, can be utilized. However,
at present, the major bottleneck for synthetic biology-based
studies on fungus/yeasts is the lack of understanding of the
post-translational modifications that strongly influence the
enzyme activity of these microorganisms. On this note, in
silico methods that help us to understand the 3D structures
of the enzymes, their interaction with the substrate, and their
Advanced Composites and Hybrid Materials (2024) 7:27
validation on the study of catalytic activity using computer
stimulation approaches might be helpful. For example, in
silico analysis of lignin peroxidase of the fungus Macropho-
mina phaseolina showed that lignin peroxidase in this fun-
gus, that had a efficient lignin degradation capacity, has a
structural similarity with the similar enzyme found in other
fungi [207].
As a concluding remark, as discussed in this review, con-
version of lignin to value-added products will be a resolution
to overcome the deep-rooted belief that “lignin can be used
to make anything but not money.” In-depth understanding
of lignin chemistry will be crucial for the development of
value-added products from it. Strategic operations can then
be followed for customized product development through
different genetic manipulation methods coupled with com-
putational applications.
Acknowledgements The authors are grateful to the authorities of
Maharaja Sriram Chandra Bhanja Deo University, Baripada, Odisha,
for providing facilities to conduct this work.
Author contribution SR, SM, and HNT contributed in drafting the
original review article and formatting. SR and DP edited the figures of
the article. HD, SM, and HNT supervised the whole work, conceptual-
ized the research, and guided the writing and editing of the article. All
authors reviewed the manuscript.
Data availability All data generated or analyzed during this study are
included in this article.
Declarations
Ethics approval and consent to participate Not applicable.
Consent for publication Not applicable.
Competing interests The authors declare no competing interests.
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