Z. Naturforsch.

2021; aop

Galena V. Angelova, Mariya S. Brazkova* and Albert I. Krastanov

Renewable mycelium based composite –

sustainable approach for lignocellulose waste
recovery and alternative to synthetic materials – a
review
https://doi.org/10.1515/znc-2021-0040 Keywords: Agaricomycetes; lignocellulose waste; myce-
Received February 11, 2021; accepted June 16, 2021; lium based composite.
published online July 12, 2021

Abstract: The agricultural waste with lignocellulose

origin is considered to be one of the major environmental
1 Introduction
pollutants which, because of their high nutritional
Development of new materials with innovative proper-
value, represent an extremely rich resource with signif-
ties is attractive trend based on scientific advances in
icant potential for the production of value added bio-
nano/materials technology because of their ability to
products. This review discusses the applications of
create matter at the atomic level. However, the process of
higher fungi to upcycle abundant agricultural by-
reproduction of live organisms is still absolutely unique
products into more sustainable materials and to pro-
and impossible to be recreated by engineering methods
mote the transition to a circular economy. It focuses on
[1–3].
the main factors influencing the properties and appli-
The introduction of living biological systems in mate-
cation of mycelium composites – the feedstock, the
rials science and nanotechnology is a strategy that has
basidiomycete species and their interaction with the
attracted considerable research efforts in recent years
feedstock. During controlled solid state cultivation on
[4–6]. This corresponds with the increasing demand for so-
various lignocellulose substrates, the basidiomycetes of
called “green” materials and production processes, which
class Agaricomycetes colonize their surfaces and form a
shifts the focus of the scientific community towards the
three-dimensional mycelium net. Fungal mycelium se-
development of new biodegradable, sustainable bio-
cretes enzymes that break down lignocellulose over time
materials derived from biological resources, as an envi-
and are partially replaced by mycelium. The mycelium
ronmentally friendly alternative to nonrenewable syn-
adheres to the residual undegraded substrates resulting
thetic materials and for decreasing pollution on the planet
in the formation of a high-mechanical-strength bio-
[7, 8]. This approach, also known as “Growing Design” [5, 9,
material called a mycelium based bio-composite. The
10], refers to growing materials from living organisms to
mycelium based bio-composites are completely natural,
achieve unique material functions, expressions and sus-
biodegradable and can be composted after their cycle of
tainable solutions for design, architecture, and industry
use is completed. The physicochemical, mechanical, and
[11].
thermodynamic characteristics of mycelium based bio-
The accumulation of large quantities of agricul-
composites are competitive with those of synthetic
tural waste with lignocellulose origin, as a result of hu-
polymers and allow them to be successfully used in the
man activity, is perceived as a serious environmental
construction, architecture, and other industries.
problem. These wastes are considered to be one of the
major environmental pollutants which simultaneously,
because of their high nutritional value, represent an
extremely rich resource with significant potential for the
*Corresponding author: Mariya S. Brazkova, Department of production of renewable energy and value added bio-
Biotechnology, University of Food Technology, 26 Maritza Blvd, materials [12–14].
Plovdiv, Bulgaria, E-mail: [email protected]. https://orcid.
In the last decade, the higher fungi of class Agar-
org/0000-0002-0184-9511
Galena V. Angelova and Albert I. Krastanov, Department of
icomycetes are in the focus of significant scientific, academic
Biotechnology, University of Food Technology, 26 Maritza Blvd, and commercial interest because of their ability to convert
Plovdiv, Bulgaria agricultural waste of lignocellulose origin into economically
2 G.V. Angelova et al.: Review of mycelium based composite production and benefits

viable and environmentally sustainable biodegradable sustainable valuable materials and to support the transi-
bio-composite materials based on the fungal mycelium tion to a circular economy.
[15, 16]. It focuses on the main factors having an effect on the
Many studies have reported the ability of basidiomy- properties and application of mycelium composites – the
cetes Lentinula edodes, Trametes versicolor, Ceriporiopsis feedstock as base, the Basidiomycetes species as matrix
subvermispora, Ganoderma lucidum, Pleurotus eryngii, and the interaction between the basidiomycetes and the
Pleurotus ostreatus and others to fully degrade various feedstock, which determines the material properties.
lignocellulose wastes [17, 18]. As a result of their vegetative
growth under controlled solid state cultivation on these
substrates, a mycelium net is formed, consisting of
numerous branched hyphae. The hyphae colonize the
2 Lignocellulose wastes as
surfaces of lignocellulose materials, decompose them promising feedstock for new
thanks to a specific enzymatic system and secretе metab-
olites that act as a natural adhesives. Fully colonized
alternative value-added bio-
lignocellulose material is heated under controlled tem- materials
perature to kill the fungal strain, to make the material inert
and to eliminate the residual water. This procedure con- Agriculture and its related industries are ones of the largest
cludes in the formation of a biodegradable, selfgrowing, sectors with the highest lignocellulose waste biomass
fibrous material with controlled physical properties, called production [22], which have special importance for the
a mycelium-based bio-composite. The mycelium-based development of bio-economy [23]. Currently, the recovery
bio-composites are completely natural and can be com- of lignocellulose residues represents a good strategy to
posted after the end of their cycle of use, which would help prevent depletion of fossil fuels, way of minimization of
the transition to a circular economy, keeping the value of greenhouse gas emissions [24] and generally add value to
materials and resources in the economy as long as possible plant biomass with lignocellulose origin [25, 26].
and generating minimal waste [3, 16, 19]. The global annual production of lignocellulose resi-
The development of composite biomaterials based on dues is in the order of 140 Gt, originated mainly by agri-
fungal mycelium, which properties are controlled and culture and agriculture-related industries [27, 28].
regulated during their formation and which would be ready Regrettably still in some countries, most of lignocellulose
for use without the need for expensive and complex pro- waste materials are left in the field to decompose or are
cessing methods, is a good strategy from economic and burned, resulting not only in significant environmental
ecological point of view [3]. Renewable mycelium-based impacts but also in the loss of potentially valuable material
composites have the potential to contribute to the new that can be used in different industries [29]. The most
economy by replacing oil-based products and converting widespread waste biomass results from different agricul-
lignocellulose waste into value-added biodegradable prod- tural and related activities – cereal husks, wheat straw,
ucts which do not damage ecosystem cycles [20, 21]. sawdust, crop stalks, dry branches, leaves, roots, fruit
Obtaining biodegradable materials that are environmen- peels, and many others [30]. Illegal lignocellulose waste
tally and socially responsible is an attractive alternative to discarding could contaminate natural water ecosystems,
using synthetic materials. The low cost, minor carbon foot- provoking process of eutrophication, together with stimu-
print and environmental impact are some of the key signif- lation of soil microbiota growth and finally induction of
icances for the use of mycelium composites as well as their harmful greenhouse gas emission possessing higher global
reduced energy consumption and biodegradability [19]. warming capacity even than CO2 [26, 30–32].
The physicochemical, mechanical and thermody- Lignocellulose wastes are considered to be one of the
namic characteristics of mycelium based bio-composites major environmental pollutants, but at the same time because
are competitive with synthetic polymers and allow them to of their high nutritional value they are accepted as globally
be successfully used in the construction, architecture and available and sustainable feedstock containing sugar-rich
design, textiles, packaging and other industries, where platform and can potentially be converted into a variety of
they can replace synthetic foams, timber and plastics [7]. different valuable products such as bio-fuels, enzymes, ani-
This review discusses the applications of higher fungi mal feed, fine chemicals, and innovative composites for civil
to recover abundant agricultural waste products into more engineering (Figure 1) [13, 14, 16, 25, 26, 30, 33–35].
 G.V. Angelova et al.: Review of mycelium based composite production and benefits 3

(mainly mannose, less glucose, and galactose), and uronic

acids. The hemicelluloses have heterogeneous composi-
tion with lower polymerization rate than cellulose and
different reactivity. The structure of hemicellulose is
mainly amorphous and more easily degradable than cel-
lulose. The main components of the hemicelluloses in
agro-agricultural waste are xylanes and mannans [41].
Lignin is one of the main components of lignocellulose
materials, which makes them resistant to physical and
biological treatments. This three-dimensional polymer
provides mainly plant resistance to insects and various
pathogens [39, 42]. Lignin structure is made up of three
phenylpropanoid units (coniferyl alcohol, sinapylalcohol,
and p-coumaryl alcohol) [43]. It is further linked to both
hemicelluloses and cellulose, forming a physical seal
around these two components. Lignin is the main barrier in
the process of degradation of lignocellulose materials and
is the most difficult component to degrade that principally
limits its application as a nutrient source and different in-
dustrial needs. Its degradation through the use of chemical
Figure 1: Options for bio-conversion of biomass into value-added and physical methods are processes that are neither envi-
bio-products [25].
ronmentally friendly, nor economically feasible [29, 36].
Many microorganisms can break down and use cellu-
Lignocellulose waste is composed of three main com- lose and hemicellulose as carbon and energy sources. But
ponents: 40–50% cellulose, 20–30% hemicellulose and only a very small number of mushroom kingdom repre-
10–25% lignin. These three components form a complex of sentatives have the unique ability to degrade lignin to CO2
nonuniform three-dimensional structures determined by [29]. These are known as wood-decay white and brown rot
the origin of lignocellulose biomass. Depending on the fungi, belonging to class Agaricomycetes. Obviously, these
source, ash, protein and pectin can also be found in the higher fungi because of their lignocellulose activity have
lignocellulose material [36]. The composition of lignocel- significant role for degradation of plant lignocellulose
lulose biomass has evolved to defy deconstruction due waste in nature [26, 35]. The ability of basidiomycetes to
to the crystallinity of cellulose, the hydrophobicity of degrade lignocellulose biomass is due to the strong
lignin, and the encapsulation of cellulose in a lignin- oxidative activity and low substrate specificity of their
hemicellulose matrix [37–39]. ligninolytic enzyme systems. The basidiomycetes are
Cellulose is the major ingredient in all plant cells and is perceived as the main recyclers on our planet. Their ability
considered to be one of the most wide spread organic mol- to break down complex organic molecules into simpler
ecules on Earth. It is a linear polymer which monomer units forms helps ecosystems to regenerate. Defeating the lignin
are anhydro-glucopyranose linked by β-1,4-glycosidic barriers is the key to the successful application of ligno-
bonds. The average cellulose chain has a degree of poly- cellulose in industrial biotechnology [29, 40, 44, 45].
merization of about 9000 to 10,000 units. Adjacent cellulose Currently, numerous research teams explore the utili-
chains bind to each other through hydrogen bonds, hydro- zation of plant lignocellulose biomass and different ba-
phobic interactions and Van der Waals forces. This leads to sidiomycetes to obtain various functional mycelium based
the parallel alignment of crystal structures and the formation composite materials to replace synthetic products
of micro fibrils [39]. More than 60% of the cellulose structure (Figure 2).
is highly organized and is treated as crystalline solid which is Lignocellulose substrates are the reinforcement of the
not accessible to water or other solvents. The close associa- mycelium-based composites and basidiomycetes are the
tion between cellulose, hemicellulose, and lignin has pro- matrix. The matrix grows on the reinforcement and fixes it
tective function against cellulose degradation [40]. in the proper position, while the reinforcement determines
Hemicellulose is the second most common component the required physical characteristics.
in the lignocellulose biomass. It is a heteropolymer of Lignocellulose waste derived from cotton, flax, hemp,
pentoses (including xylose and arabinose), hexoses rice, sorghum, and wheat are the most studied substrates
4 G.V. Angelova et al.: Review of mycelium based composite production and benefits

Figure 2: A mycelium-based composite con-

sists of a lignocellulose reinforced fungal
mycelium.

for growing fungi that enhance structural properties and more ‘circular’ economy supporting sustainable food pro-
result in obtaining new mycelium-based composites [3, 12, duction, manufacturing and energy production, and new
46–49]. Coffee grounds, pine waste, and general wood materials obtaining [15, 54–58].
waste are chosen as a substrate by Alves et al. [50]. The Studies on the use of higher fungi to derive new bio-
properties of the mycelium composites are strongly materials began in the 1990s by Japanese scientist Shigeru
affected by the type of the substrate. According to Elsacker Yamanaka, who used mushroom mycelium for the pro-
et al. [51], the substrate fiber condition (loose, chopped, duction of paper and building materials [59]. In 2007 in the
dust, precompressed, and tow), size, and processing in- US Eben Bayer and Gavin McIntyre established the Eco-
fluence the characteristics of the bio-composites more vative Company. They, together with the scientist Philip
strongly than the chemical composition. Haneef et al. [3] Ross are leaders in R&D of mycelium based materials.
concluded in his research that the obtained fibrous myce- Research in Europe on mycelium-based composites has
lium material is mechanically more rigid when fungus are been initiated by Mauricio Montalti in collaboration with
grown on amorphous cellulose and it is harder to digest the University of Utrecht (the Netherlands). They are
compared to the cellulose substrate containing potato exploring the possibility of producing mycelium-based
dextrose broth (PDB)). The selection of appropriate ligno- composites as tests using local waste streams of the Eu-
cellulose substrates suitable for fungal growth and ropean agricultural industry. These experiments were first
mycelium-based material obtaining is a key step in the funded by the NLF (Stimuleringsfonds) and later by na-
development of an applicable technology. tional and European research institutions.
Fungi are a large group of diverse unicellular, multi-
cellular or syntactical spore-producing organisms which
feed on organic matter [19]. Often referred to as the
3 Basidiomycetes as a matrix for “Forgotten Kingdom”, fungi are key components of life on
new mycelium-based materials this planet [60]. Fungal diversity is thought to exceed that
of terrestrial plants at a ratio of 10:1 [61].The phylum
The most common processes of bio-fabrication mainly Basidiomycota is considered the best choice for mycelium
focus on microbial transformation of the lignocellulose composite development, among the fungi, due to their
biomass feedstock to monosugars by enzymatic hydrolysis ability to form more complex and larger organic structures
followed by conversion to higher-value products. This compared to other fungal representatives [45].
strategy requires resource-intensive biomass processing, The subkingdom Dikarya incorporates the extraordi-
and special equipment [52, 53]. On the other hand, higher narily diverse and large phylum of basidiomycetes,
fungi could grow directly on lignocellulose substrates and together with the ascomycetes. Most basidiomycete species
could assure simpler, less resource-intensive means of belong to the filamentous fungi. They have complex life-
production and at the same time to expand the possibilities cycles, reproduce sexually and produce basidiospores on
for the production of bio-materials [36, 54]. The controlled specialized cells called basidia. Mushrooms from class
solid state cultivation of higher fungi could also nurse a Agaricomycetes make up to 98% of the Agaricomycotina –
 G.V. Angelova et al.: Review of mycelium based composite production and benefits 5

the largest subphylum of basidiomycetes [60]. Their apical region of hyphal tip [19]. Hypha initiate random
morphogenesis is associated with the formation of branching after an isotropic growth phase, starting formation
macroscopic fruiting bodies, also known as basidioma or of fractal tree-like colonies. Colonies associate randomly
basidiocarp. These fruiting bodies are large enough to be through hypha fusion forming a random fiber network
seen with the naked eye and can be touched by hand, structure. The branching density and network topology are
which is why these mushrooms are also called macro- dependent on the nutritional and growth conditions [68, 69].
mycetes. One-third of the described species of fungi be- The filamentous mode of fungal growth stimulates effective
longs to Agaricomycetes and includes the most outstanding colonization of substrates and provides a large surface to
basidiomycete species, all of which produce basidiocarps, volume ratio facilitating the uptake of nutrients [19].
such as the gilled mushrooms, bracket fungi, puffballs, The presence of septa is an important feature in the
crust fungi, chanterelles, coral fungi, and jelly fungi [62]. production of mycelium-based materials [19, 50]. Septa,
Lignocellulose in wood substrates could be decomposed special transverse cell walls, have an opening that can be
only by a type of fungi called wood-decay fungi. A large closed in order to block the draining of a cytoplasm
number of those fungal representatives are able to degrade through the rapture when a hypha becomes damaged. This
lignocellulose thus releasing vital nutrients back to the will decrease the damage to the colony and therefore will
environment. The evolution process of fungi made possible lead to faster colonization of a substrate. As a result, the
different types of basidiomycetes to degrade specific types of presence of septa significantly increases the strength of the
organic compounds. Wood decaying fungi are generally mycelium, and hence the strength of the obtained bioma-
divided into two main types: brown and white rot [60]. More terial [19, 45, 68, 69].
than 90% of basidiomycetes belong to the so-called white rot The third important feature is the process of anasto-
fungi. Their abundance in nature coupled with the ability to mosis. Anastomosis is the ability of two different hyphae to
completely mineralize lignocellulose cell wall polymers and fuse when they meet. Anastomosis is crucial for the crea-
convert them into carbon dioxide and water makes these tion of fast-growing mycelium as it allows the creation of
fungi important players in the recycling of carbon in nature well-developed mycelium net and the resulting mycelium
and determines their current application in industrial mass is much more coherent [19, 45, 70].
biotechnology. Although brown rot fungi constitute only The cell wall plays some physiological roles in fungal
about 7% of wood rotting basidiomycete fungi, they are also morphogenesis (Figure 3), protecting hypha against os-
valuable recyclers of lignocellulose biomass. They rapidly motic lysis and providing mechanical strength to the whole
depolymerize cellulosic materials while only modifying mycelium [3, 71]. Chitin, glucans, manoproteins, chitosan,
lignin via a Fenton reaction based mechanism [40]. polyglucuronic acid, cellulose, and specific proteins and
Basidiomycetes possess some important characteris- glycoproteins are the major compounds in the cell wall and
tics that make them preferable for the production of are thought to have a positive effect on the mechanical
mycelium-based materials – filamentous mode of growth, strength and thermal stability of the obtained mycelium-
the presence of septa, the process of anastomosis, the based composites [3, 68].
structure, and compounds of the fungal cell wall and Chitin is a linear polymer of the acetylated amino sugar
specific enzymatic systems. N-acetylglucosamine which is very strong and has a tensile
The filamentous mode of growth of higher fungi as a new strength significantly greater than many synthetic mate-
form of low energy bio-production and waste recovery is in rials such as carbon fibers and steel due to hydrogen
the focus of academic and commercial community over the bonding along the chains which give them rigidity [19, 20,
past decade [19, 63–66]. The vegetative lower part of mac- 72].The microfibers of chitin are responsible for the firm-
romycetes constitutes a network of fine white mycelium. ness and mechanical strength of the hyphae, while the
Mycelium grows due to symbiotic relationships with the behavior of the mycelium under mechanical stresses is
nutrient substrates. By using physical pressure and enzyme characterized by elastic properties, orientation, and con-
systems the mycelium covers or penetrates the substrate. The nections of the hyphae [50, 68]. Hydrophobins are low-
secreted enzymes break down the polymers into easily weight proteins found in fungi which act by selfassembling
absorbed and transported sugars [3, 67]. The porous struc- to form an amphipathic membrane due to their hydro-
ture of the mycelium is composed of tubular filaments called phobic nature. Hydrophobins are involved in a number of
hypha. Typically, hyphae have diameters in the range of 1– functions related to cell wall morphogenesis, hydropho-
30 μm, which could vary depending on the species and bicity, and substrate adhesion, both in aqueous and aerial
growth conditions, and lengths ranging from a few microns environments and favor the thermal stability of mycelium-
to several meters [50, 68]. Mycelium extension occurs in the based material [70, 73, 74].
6 G.V. Angelova et al.: Review of mycelium based composite production and benefits

Figure 3: Representation of mycelium

physiology [3].

Higher basidiomycetes are predominantly responsible The patent of Ecovative Design LLC uses P. ostreatus,
for lignocellulose degradation and they are the most rapid Fomes fomentarius, Ganoderma aplantum, and Inonotus
lignocellulose degraders. That is why basidiomycetes are obliquus and wheat hulls or cotton seeds for a fungi-
of fundamental importance for the efficient bioconversion derived biofoams production process [84].
of lignocellulose residues and they are a promising agent in Pleurotus pulmonarius, P. ostreatus, Pleurotus salmo-
obtaining mycelium based materials [7, 19, 27, 44, 75–77]. neostramineus, and Aaegerita agrocibe were grown on
The efficiency of lignocellulose degradation could be woodchips of eucalyptus, oak, pine, apple, and vine in
associated with a mycelium growth manner that allows the order to evaluate which pair of fungi-plant material pair
fungus to transport scarce nutrients such as nitrogen and was the most suitable combination for future applications
iron, to a distance into the nutrient-poor lignocellulose for producing mycelium based composites. The research
substrate that constitutes its carbon source [78]. shows that the most efficient integrations are the samples
The fungal degradation occurs exocellularly, either in of P. ostreatus grown on apple or vine woodchips [47].
association with the external cell membrane layer or Twenty-seven different fungal species were used by
extracellularly, because of the insolubility of lignin, cel- Cerimi et al. [66] in the developing patent for mycelium
lulose and hemicellulose. Fungi possess two types of based materials and processes. Mycelia from two types of
extracellular enzymatic systems: the hydrolytic system and edible, medicinal fungi G. lucidum and P. ostreatus fed by
a unique oxidative extracellular ligninolytic system. The two bio-substrates were studied by Haneef et al. [3].
hydrolytic system consists of hydrolases that are respon- Depending on the substrate the obtained fibrous
sible for polysaccharide degradation while the ligninolytic composites show different morphology and mechanical
system degrades lignin and opens phenyl rings. White rot properties. Sawdust and wheat bran were used by Bruscato
fungi, as primary decomposers, break down cellulose and et al. [48] as a feeding substrate in a controlled cultivation
hemicellulose mainly through hydrolytic enzymes such as of Pycnoporus sanguineus, Pleurotus albidus and Lentinula
cellulases and xylanase, and lignin is degraded by oxida- velutinus in order to receive biofoams to replace expanded
tive ligninolytic enzymes such as lignin peroxidase (LiP) polystyrene. T. versicolor and five different types of sub-
and manganese peroxidase (MnP). The use of basidiomy- strate (hemp, flax, flax waste, softwood, straw) and sub-
cetes in the processes of digestion of lignocellulose wastes strate conditions (loose, chopped, dust, precompressed,
is considered an attractive alternative for their conversion and tow) were investigated in order to elucidate the pro-
into biological value products. duction process, the mechanical, hygrothermal, and
Many studies have reported the ability of L. edodes, chemical properties of formed mycelium-based composites
T. versicolor, C. subvermispora, G. lucidum, P. eryngii, P. [51].
ostreatus and others to be cultivated on a range of lignocel- Agaricus bisporus, Lentinula, P. ostreatus, G. lucidum,
lulose materials, including various types of cotton seed hulls, Pleurotus ostreatussajorcaju, Pleurotus ostreatusflorida,
corn cobs, peanut shells, cotton from the textile industry, Kuehneromyces mutabilis, and Flammulina velutipes were
coffee pulp, paper [29], and leaves [79] as well as low-quality cultivated on oat husk and rapeseed cake under controlled
organic waste streams like saw dust and straw [15, 18, 80]. growth conditions and the novel mycelium composites was
The effect of the choice of fungal strain and the sub- characterized by Tacer–Caba et al. [85]. The results
strate composition on the macroscopic properties of demonstrated that the mycelium composites were hydro-
mycelium-based materials is the subject of research of phobic and robust when grown on rapeseed cake. A. bis-
many recent studies [3, 19, 20, 47, 68, 77, 81–83]. porus grown on rapeseed cake exhibited increased stiffness
 G.V. Angelova et al.: Review of mycelium based composite production and benefits 7

after humidity was successively increased and decreased. The production process of mycelium-based products is
The moisture-resistance of these novel mycelium com- completely different from the traditional industrial pro-
posites is encouraging for novel sustainable material cesses. The introduction of a living biological system in the
solutions. first stages of processing is an innovative strategy that has
attracted considerable research efforts in recent years [4,
6].
4 Mycelium based composites as The process starts with solid-state cultivation of
selected fungal strains on previously chosen and appro-
an alternative to synthetic priately treated lignocellulose waste substrate or a mix of
materials substrates. The adequate sterilization of the substrates has
an important role as a prevention measure against unde-
Synthetic materials and products derived from petroleum sirable microbial contaminations together with the
derivatives have been widely used in human life and in obtaining of acceptable results in the process of mycelium
various industries [86, 87]. Their production is adversely based composites production [19]. The cultivation param-
affected by resource depletion, and due to the fact that they eters (light, temperature, and humidity) should be accu-
are not biodegradable, they cause serious environmental rately maintained for about 20 days to allow stable growth
damages. Improper and uncontrolled disposal of synthetic of the basidiomycetes [105]. The optimal conditions of
polymeric materials also compromises natural biodegra- temperature and moisture vary considerably depending on
dation processes, as some compounds present in synthetic the strain of fungi employed. The optimal temperature for
polymer chains can act as a barrier against the degradation satisfactory growth is between 25 and 35 °C and the pref-
of organic compounds [88]. erable humidity is around 60–65% [19]. The humidity value
This directs research to seek new approaches for the is essential for fungal metabolism and prevents of sub-
production of polymeric materials based on renewable strate from drying [105]. The formation of fruiting bodies as
resources [89–91]. Intensive research efforts focus on the well as the obtaining of rapid mycelium growth is accom-
development of polymeric materials of natural plant, ani- plished by cultivation in darkness [11, 106]. During culti-
mal and bacterial origin – plant cellulose [92], lignin [93], vation the fungal strain degrades the lignocellulose
pectin [94, 95], plant and animal proteins [96, 97], bacterial substrates and gradually covers it with a three-dimensional
polyesters [98], or plants [99]. They are all durable mate- mycelium network. The mycelium secretes enzymes that
rials, biocompatible and biodegradable with a wide variety break down lignocellulose polymers into substrates that
of properties. However, the fabrication of such bio- are used as nutrients. As a result, the organic substrates are
polymers involves the application of complex, costly, and degraded over time, being fully or partially replaced by
inefficient methods of processing their bio-sources for their fungal biomass, and the accumulated mycelium strongly
extraction and functionalization [100]. For this reason, adheres to the residual lignocellulose and forms the
although these materials can solve various environmental structure core of the mycelium composites. To acquire
problems, they are still expensive and have very limited products with desired geometry different molds could be
application possibilities [101, 102]. used. Different 3D printing technologies could also be used
The development of mycelium based bio-materials in order to overcome differences of molds using [50]. The
i.e., mycelium composites and pure mycelium, based on production of mycelium-based composites includes inac-
fungal mycelium, which properties are controlled and tivation of the fungus by heat treatment at a minimum of
regulated during their formation and which would be ready 60 °C at some point during colonization [107]. The time
for use without the need of expensive and complex pro- needed for colonization before inactivation determines the
cessing methods, is a good strategy for overcoming these ratio between lignocellulose substrate and fungal biomass
problems [3, 103]. [15, 50]. Pure mycelium material is obtained by complete
Renewable mycelium-based materials have the po- degradation of the lignocellulose by fungus and its
tential to contribute to the new economy by replacing oil- replacement with mycelium. The duration of the process
based products and converting lignocellulose wastes into varies with the species, substrate, growth conditions, and
value-added products that are biodegradable and do not application as a final product [3, 68, 81, 108]. It is clear that
damage ecosystem cycles. The obtaining of biodegradable full degradation of the substrate is time consuming and the
materials that are environmentally and socially respon- outcome is a larger weight loss of the material, compared to
sible is an attractive alternative to using synthetic materials the breakdown needed to produce mycelium composites.
[3, 19, 46, 82, 104]. Pure mycelia are more expensive to produce [15, 50].
8 G.V. Angelova et al.: Review of mycelium based composite production and benefits

The properties of the composite are affected not only synthetic polymers and engineered woods for insulation,
by the used species, but also by the substrate and the furniture, and panels applications [21].
growth conditions. When those parameters are constant A non-traditional technique for the obtaining of bio-
the result would be a mycelium composite with determined composites based on natural textile fiber, mycelium-bound
compressive strength, electrical conductivity, and flexi- agricultural waste, and bio-resin matrix was explored by
bility [19]. Jiang et al. [112]. The study showed that the type of sub-
According to some recent studies, mycelium-based strate, degree of colonization within the composite surface
bio-composites have characteristics that are competitive by mycelium, and bonding between the core and surface
with synthetic polymers (Figure 4). Mycelium-based bio- strongly affected the strength of the panel. The rigidity was
composites have low density and good acoustic and ther- dominated by core strength depending on its thickness.
mal insulation properties [3, 7, 70, 103, 109]. Six blends of The physicochemical, mechanical and thermodynamic
processed cotton plant biomass materials were developed characteristics of the mycelium-based bio-composites allow
as a substrate for selected fungi and assessed by Holt et al. them to be successfully used in the construction, architec-
[63] for the production of packaging material based on ture, and design, textile, packaging and other industries for
molds. They stated that the developed bio-composite was a non-structural (insulation and door cores), semi-structural
valuable alternative to polystyrene foam packaging mate- (paneling, flooring, cabinetry, and furnishing), and specific
rial and had similar mechanical properties. applications [3, 70, 103, 109, 112–115]. Foam-like mycelium
The thermal properties of mycelium based insulation bio-composites due to their thermodynamic properties and
materials for construction were evaluated by Xing et al. acoustic absorbance show great potential for thermal and
[110] and they concluded that the materials demonstrated acoustic insulation in construction industry, packaging,
good thermal performance. Travaglini et al. [111] suggested and infrastructure development [116]. Oriented strand board
that mycelium foams could provide eligible mechanical (OSB) and medium density fiber board (MDF), that often
properties and also were lightweight, sustainable and contain formaldehyde resin could be replaced by nontoxic
inert. The compressive strength of the material was almost natural-like mycelium bio-composites with decreased water
three times as high as its tensile strength. The mycelium absorbance, mechanical and physical properties, suitable
bio-composite is closest to polystyrene foam based on its for indoor applications [83, 117]. Some textiles and leathers
density and strength properties. A number of mycelium- with significant carbon footprint are considered to be fully
based composite materials were tested for their fire safety. replaceable by pure mycelium materials thus leading to
It was found that the bio-composites were a very good obtaining lower or even negative footprint [114]. Many artists
alternative to highly flammable petroleum-derived and designers are interested in mycelium-based materials
for production of furniture, lampshades and haute couture
clothes, bags, and shoes [7, 118]. Lakk et al. [119] investi-
gated the production process and feasibility of fungal based
bio-composite material for habitat structures on the Moon
and Mars, using automated additive construction technol-
ogy. Most of the current research of NASA Research Center in
California focuses on developing and testing prototype
technologies that may enable future explorers to ‘grow’ their
own houses for use in Moon and Mars colonies. The system
would involve the mycelia produced by white rot fungi.
These technologies also have the potential for creating
sustainable structures on Earth [120, 121].

5 Conclusion
Lignocellulose waste originated from agriculture and its
related industries is especially plentiful in nature and has
an enormous potential for bioconversion. Agglomeration
of lignocellulose in large quantities, in places where agri-
Figure 4: Comparison polystyrene and mycelium foams [19]. cultural residues present a disposal problem, results not
 G.V. Angelova et al.: Review of mycelium based composite production and benefits 9

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