9 - Paper 9 - Original

microorganisms
Review
Using Fungi in Artificial Microbial Consortia to Solve
Bioremediation Problems
Elena Efremenko * , Nikolay Stepanov , Olga Senko , Aysel Aslanli , Olga Maslova and Ilya Lyagin
Faculty of Chemistry, Lomonosov Moscow State University, Lenin Hills 1/3, Moscow 119991, Russia;
[email protected] (N.S.)
* Correspondence: [email protected]; Tel.: +7-(495)-939-3170; Fax: +7-(495)-939-5417
Abstract: There is currently growing interest in the creation of artificial microbial consortia, especially
in the field of developing and applying various bioremediation processes. Heavy metals, dyes,
synthetic polymers (microplastics), pesticides, polycyclic aromatic hydrocarbons and pharmaceutical
agents are among the pollutants that have been mainly targeted by bioremediation based on various
consortia containing fungi (mycelial types and yeasts). Such consortia can be designed both for the
treatment of soil and water. This review is aimed at analyzing the recent achievements in the research
of the artificial microbial consortia that are useful for environmental and bioremediation technologies,
where various fungal cells are applied. The main tendencies in the formation of certain microbial
combinations, and preferences in their forms for usage (suspended or immobilized), are evaluated
using current publications, and the place of genetically modified cells in artificial consortia with fungi
is assessed. The effect of multicomponence of the artificial consortia containing various fungal cells is
estimated, as well as the influence of this factor on the functioning efficiency of the consortia and the
pollutant removal efficacy. The conclusions of the review can be useful for the development of new
mixed microbial biocatalysts and eco-compatible remediation processes that implement fungal cells.
Keywords: consortium composition; main trends; genetically modified cells; multicomponence effect;
pollutant removal efficacy; immobilized forms
Citation: Efremenko, E.; Stepanov, N.;

Senko, O.; Aslanli, A.; Maslova, O.;
Lyagin, I. Using Fungi in Artificial 1. Introduction
Microbial Consortia to Solve
Bioremediation processes are becoming increasingly important, both from the point
Bioremediation Problems.
of view of developing the scientific basis for their implementation, and the point of
Microorganisms 2024, 12, 470.
https://doi.org/10.3390/
view of their practical application [1]. Soil and water ecosystems contaminated with
microorganisms12030470
oil and petroleum products [2], heavy metals [3], polycyclic aromatic compounds [4], and
pesticides [5] have been mainly referenced in discussions of the objects of bioremediation.
Academic Editor: Giovanni Vallini Now these pollutants [6] and microplastics [7] have been joined by pharmaceutical agents.
Received: 21 January 2024 The biodegrading microorganisms of various pollutants have recently generated partic-
Revised: 12 February 2024 ular interest in studies of consortia of microorganism cells with heterogeneous composition,
Accepted: 23 February 2024 which combine the different metabolic potentials of participants [8–10]. In most cases,
Published: 26 February 2024 natural consortia contain a significant number of different prokaryotic cells (bacteria and
archaea) that form biofilms, which may include aerobic and anaerobic cells [11]. At the
same time, the extremely interesting properties possessed by those variants of consortia
are created by cells of these prokaryotic microorganisms (i.e., bacteria and archaea) and
Copyright: © 2024 by the authors. eukaryotic microorganisms (in particular, filamentous fungi and yeast), which are able
Licensee MDPI, Basel, Switzerland. to coexist and catalyze biodegradation and bioremediation processes [12]. Such an arti-
This article is an open access article ficial consortia, involving cells of microorganisms from different kingdoms [13–17], are
distributed under the terms and
especially attractive from a scientific and practical point of view, due to the possibility
conditions of the Creative Commons
of expanding its total metabolic potential. At the same time, all the advantages that the
Attribution (CC BY) license (https://
artificial consortia of microorganisms themselves have over natural analogues, which have
creativecommons.org/licenses/by/
mainly been described in the literature on bacterial cells, are preserved:
4.0/).
Microorganisms 2024, 12, 470. https://doi.org/10.3390/microorganisms12030470 https://www.mdpi.com/journal/microorganisms

Microorganisms 2024, 12, 470 2 of 26
- easy-to-use reproducibility of the compositions of the consortia;

- the possibility of introducing maximum targeted metabolic activity into the consortia
cells, which are improved, including through the genetic modification of cells;
- targeted variation of the ratios of cell concentrations in the consortium, to regulate the
rates of associated biochemical processes catalyzed by cells;
- sustainable functioning of the created consortia, which are accompanied by increased
degrees of conversion of the original substrates, including toxic compounds [18–26].
The growing interest in fungi as part of artificial consortia is due to the fact that fungi
are capable of synthesizing a very wide range of different hydrolytic and redox enzymes.
A large number of pollutants can serve as substrates for these enzymes and undergo
deep conversion due to their broad substrate specificity [12,17]. In addition, most of these
enzymes are secreted and catalyze the destruction of substrates outside the cells of the fungi
themselves, which provides more easily accessible sources of nutrition to all participants in
the formed consortia. Moreover, fungi, being part of such consortia, are able to function in
a wide range of pH and temperature values, in aerobic and anaerobic conditions [27–29].
In addition to the various catalytic capabilities that are characteristic of fungi, it has
been noted that the morphology and biochemical composition of yeast cell walls, as well as
the hyphae of filamentous fungi, can help reduce the level of toxicity of consortium habitats
in the presence of various pollutants, including heavy metals, due to their biosorption [30].
Another mechanism for reducing the toxicity of media, which can be implemented by
fungi in consortia with their participation in bioremediation processes, is based on their
synthesis of lipopeptide biosurfactants. The appearance of these group of biosurfactants
among metabolites can lead to biodeposition of pollutants [27]. In turn, the accumulation
of organic acids, actively produced by fungi [31,32], as fungal metabolites in environments
with functioning consortia of microorganisms promotes the formation of complexes with
metals that prevent the migration of these pollutants.
Thus, the information that has been accumulated today indicates that the bioreme-
diation of various environments, with the participation of fungal cultures as part of the
used consortia, is implemented using biosorption, complexing and other physical and
chemical processes, in addition to extracellular catalytic and intracellular metabolic re-
actions. Moreover, compared with the design of bacterial artificial consortia, in order
to solve similar issues of bioremediation in synthetically formed biosystems containing
fungal and yeast cultures, it is necessary to not only take into account the compatibility and
antagonism of different microorganisms, but to also consider the features of cross-kingdom
reactions [16,20]. In particular, chemically similar quorum-sensing signaling molecules
such as lactone-containing molecules can appear in the microenvironment of the cells of dif-
ferent microorganisms (bacteria and fungi) which can, in turn, contribute to the formation
of a synergistic effect from their participation in consortia.
On the other hand, it is known that fungi in a quorum state are capable of synthesizing
antibiotics and mycotoxins, which limit their use in artificially formed consortia [33].
The purpose of this review is to analyze and compare the information accumulated
in recent years on artificial consortia of different compositions with the obligatory par-
ticipation of fungal cells in them. This analysis aimed to highlight the main priorities in
modern experimental developments carried out by different researchers in the creation and
application of mixed artificial consortia, and to also try to highlight current trends in the
development of scientific and practical information in the field that is developing artificial
mixed consortia with the participation of fungal cells.
2. Main Targets for Bioremediation Based on Various Consortia Containing Fungi

Recently, much attention has been paid to discussing the principles of designing
artificial consortia [1,18,20,34–36]. In this review it was decided to analyze the success of
already obtained consortia in the processes of bioremediation of various contaminated
environments, taking into account the composition of microorganisms introduced into the
consortia. In this regard, special attention was paid to exactly which microorganisms were
used in consortia with fungal cultures to remove specific pollutants (heavy metals, dyes,
synthetic polymers, pesticides, polycyclic aromatic hydrocarbons, pharmaceutical agents,
crude oil, phenols, etc.), and to the degree that their removal was achieved.
2.1. Removal of Heavy Metals

Heavy metals which, as a result of anthropogenic pollution, enter the environment
and accumulate in food chains, have a negative effect on the state of the cells of various
living organisms [37]. Lead, cadmium, and chromium are highly toxic, mutagenic, and
carcinogenic, and the presence of several metals in contaminated objects at once can
aggravate their toxicity. Metal ions can interact with hydroxyl, carboxyl, phosphate and
sulfhydryl groups, causing changes in the properties of proteins, nucleotides, coenzymes,
phospholipids and other substances involved in cell metabolism. In addition, metal ions
can be replaced in the active centers of enzymes, causing inhibition of their activity [1].
Soil contamination with heavy metals leads to a decrease in the species diversity of
microbial communities, although the cells of individual microorganisms are capable of
accumulating heavy metals, due to their sorption and intracellular complex formation. The
most active participants in these processes are polysaccharides of the cell walls of various
microorganisms [38], especially those that are part of consortia (Table 1) [39–44].
Table 1. Microbial consortia with fungal species for removal of heavy metals.
Consortia [Reference] Conditions Pollutant/Process Efficiency

Aspergillus niveus, A. flavus, 1.4 ×106 spore/mL of each strain; Removal of Cr, Zn, Pb, Cd and
A. niger [39] pH 5.0, 110 rpm, 30 ◦ C, 96 h Ni—70–90%
Heavy metal concentration—100 mg/L, Removal of Cr(VI)—81%, Cd(II)—82%,
A. flavus, A. fumigatus [40]
1.2 × 106 spores/mL, pH 5.0, 30 ◦ C, 144 h mixture of metals—73%
Initial metal concentration
Removal of As—77%, Cr—60%,
Ascomycota and Basidiomycota fungi [41] (23–2347 mg/kg), pH 7.9,
Cu—52%, Fe—52%, Mn—71%
soil moisture 60–65%, 28 ◦ C, 100 days
Initial metal concentration (400–800
Removal efficiencies of Ni, Pb, Zn—52%,
Ascomycota and Basidiomycota fungi [42] mg/kg), pH 7.9, soil moisture 60–65%,
44%, 32% respectively
28 ◦ C, 100 days
A. fumigatus, A. terreus,
Cd—100 mg/L, pH 5.0, 30 ◦ C, 120 h Removal of Cd(II)—95%
Paenibacillus dendritiformis [43]
Pb(II)—20.5–293.23 mg/L,
A. terreus, Talaromyces islandicus,
Ni(II)—12.1–164.7 mg/L, Removal of Pb(II) and Ni(II)—95–97%
Neurospora crassa, Aspergillus flavus [44]
inoculum 8%, pH 5.0, 30 ◦ C 120 h
Bioremediation of heavy metals by microorganisms includes the following main mech-

anisms: adsorption, bioaccumulation, biotransformation and bioleaching [45]. Microbial
consortia of different compositions are characterized by different abilities to remove heavy
metals. The majority of interest is focused on those options that can be used for bioremedi-
ation of objects contaminated simultaneously with several heavy metals. For example, a
fungal consortium consisting of several species of fungi of the genus Aspergillus was used
to remove heavy metals [39].
It was found that such a consortium leads to an increase in the removal of heavy metals
Cr, Zn, Pb, Cd and Ni by 2–45%, in comparison to individual fungal cultures included
in the consortium. At the same time, the efficiency of bioremediation of heavy metals
under the influence of such a consortium reached 70–90%. Interestingly, Cr removal was
maximum in the consortium of A. niveus and A. flavus (93%), while in the combination of
A. niger with A. flavus or A. niveus cells it decreased by 3%. Thus, the results obtained from
combining fungi belonging to the same genus were slightly different.
The genetic proximity of fungi used in consortia does not lead to a significant im-
provement in their characteristics in bioremediation processes [40]. Thus, an attempt by
other researchers to increase the efficiency of removal of Cr(VI)) and Cd(II) (100 mg/L) by
a fungal consortium consisting of a strain of A. flavus and two closely related strains of
A. fumigatus only made it possible to obtain 80% for each of the metals, and slightly more
than 70% when processing a mixture of both metals by this consortium.
A fungal consortium consisting of 13 strains of various fungi (Perenniporia subtephropora,
Aspergillus fumigatus, A. niger, Phanerochaete concrescens, Cerrena aurantiopora, Polyporales
sp., Fusarium equiseti, F. chlamydosporum, Paecilomyces lilacinus, Tremates versicolor, Antrodia
serialis, Daldinia starbaeckii, Penicillium cataractum) was effective in removing metals As
(77%), Mn (71%), Cr (60%), Cu (52%) and Fe (52%) from the soil within 100 days [41].
It was observed that when treated with contaminated soil the number of fungal
cells increased until day 60, and then decreased. Most likely, this could be due to cells
accumulating a certain maximum concentration of metals, which then began to have a
negative effect on the functioning of the consortium. In support of this explanation, when
the same consortium was used in another study to remove Ni, Pb, Zn from soil, the metal
removal efficiency was 32–52%, as a result of bioaccumulation [42].
In optimal conditions (in the presence of glucose and peptone, pH 5.0, 30 ◦ C, 120 h), the
efficiency of the microbial consortium consisting of A. fumigatus, A. terreus and Paenibacillus
dendritiformis in removing Cd was 95% [43]. Thus, in this case, adding just one “outsider”
(that is, bacteria P. dendritiformis) to the consortium of filamentous fungi belonging to the
same genus led to a more significant efficiency of the process than when using only “close
relatives” in the created consortium.
In a similar situation with two representatives of the genus Aspergillus (Aspergillus
terreus, A. flavus), the introduction into a consortium of not one, but two, “outsiders”
(Talaromyces islandicus, Neurospora crassa) for the removal of Pb, Ni from real wastewater
gave a guaranteed high efficiency of simultaneous removal of both metals (95–97%) [44].
In general, known examples of the use of fungal consortia clearly show the successful
removal of heavy metals through their adsorption and accumulation in cell biomass with
higher efficiency, compared to the use of individual strains of microorganisms. At the
same time, the most attractive option is to combine a small number of participants in
such consortia, for instance combining fungi from the same genus with the addition
of “exogenous” representatives, presumably so they are playing the role of competitors
or antagonists.
2.2. Decolorization of Dyes

Various industries use dyes, including synthetic ones, with complex aromatic molec-
ular structures, whose degradation poses a serious problem [46]. Many physicochemical
processes, including adsorption, chemical oxidation, and photocatalysis, are effective in
decolorizing wastewater containing dyes. However, high cost and the formation of toxic
secondary compounds limit the use of these treatment methods. For comparison, the
biological degradation of a number of dyes, for which the use of fungi has been shown
to be highly effective, is relatively more economically profitable and environmentally
friendly [47].
Decolorization and degradation of dyes by fungi are one of the most studied processes,
accounting for hundreds of articles published in the field [48]. Fungi remove dyes through
processes such as biosorption, biodegradation, and bioaccumulation. The degradation
potential depends primarily on fungal metabolism and the expression of extracellular en-
zymes such as lignin peroxidase (LiP), manganese peroxidase (MnP) and laccase. However,
high efficiency has only been established for the decomposition of certain dyes, while
wastewater from enterprises is usually a mixture of various organic and inorganic pollu-
tants, including dyes. The use of microbial consortia may therefore be more effective for
dye removal (Table 2) [49–56].
Yeasts, as eukaryotic and single-celled fungi, also have practical applications in
wastewater treatment, due to their high degradation activity in various azo dyes. In
particular, a yeast consortium consisting of three different cultures was obtained that was
capable of biodegrading industrial dyes [49].
Table 2. Microbial consortia with fungal species for decolorization of dyes.

Yarrowia sp., Barnettozyma Degradation of Scarlet GR, Red HE3B, Remazol
100 mg/L of dye,30 ◦ C,
californica, Sterigmatomyces Brilliant Blue R, Methyl Orange, Rubine GFL and
static conditions, 6–12 h
halophilus [49] Reactive Red 2—92–100%
Daldinia concentrica, 50 mg/L of dye, pH 4.5, 30 ◦ C,
Degradation of cibacron brilliant red 3B-A—99%
Xylaria polymorpha [50] 150 rpm, 48 h.
Rhodotorula sp., Raoultella
Degradation of methylene blue—100% and
planticola and Staphylococcus 200 mg/L of methylene blue in municipal
78.5% in municipal wastewater and industrial
xylosus cells immobilized in wastewater and industrial effluent, 144 h
effluent, respectively
Ca-alginate beads [51]
Degradation of reactive blue 4, fast green, methyl
red, crystal violet, alura red AC, tartrazine,
A. niger, A. terrus, A. oryzae, 20 mg/L of each dye, 150 rpm, naphthol blue black, janus green B, alizarin
A. fumigatus [52] 28 ◦ C, 72 h yellow R, evans blue, brilliant green,
pararosaniline, ponceau S, cibacron brilliant red
3B-A, direct violet 51—57–100%
Aspergillus sp., Disperse Red—0.1 g/L, pH 6.0, Degradation/adsorption of disperse red
Chlorella sorokiniana [53] 160 rpm, 25 ◦ C, 4 days 3B—98.1%
Daedalea dickinsii, Methyl orange—100 mg/L, 30 ◦ C,
Degradation of methyl orange—98%
Pseudomonas aeruginosa [54] 7 days
Reactive Black 5, Acid Orange 7; Reactive
Green 19, Reactive Yellow, ABC, Atlantic
Sterigmatomyces halophilus,
Black C—50 mg/L, glucose as Degradation—88–97%
Meyerozyma guilliermondii [55]
co-substrate,
pH 7.0, 35 ◦ C, 120 h
Penicillium oxalicum, 100 mg/L of congo red with dextrose
Congo red degradation—97.1%
Aspergillus tubingensis [56] (10 g/L), pH 5, 150 rpm, 28 ◦ C, 12 h
It was shown that when such a yeast consortium is used, various dyes (100 mg/L) can be
degraded within 6–12 h with an efficiency of 90–100%. In real wastewater from the textile
industry, a significant reduction in the content of total organic carbon (54%), biological
oxygen demand (74%) and chemical oxygen demand (95%) and color removal (98%) was
shown within 24 h at 30 ◦ C and pH 8.0 in the same consortium. The synthesis and secretion
by cells of the consortium of a number of redox enzymes (laccase, veratryl alcohol oxidase,
tyrosinase, azoreductase, NADH-DCIP reductase), that are found in the reaction medium,
ensured the efficiency of the process [49].
The efficiency of bleaching a dye such as cibacron brilliant red 3B-A by a consortium
of fungi Daldinia concentrica and Xylaria polymorpha reached 98–99%, and was higher in
comparison with the effect of the same cultures used separately [50]. Among the enzymes
involved in dye degradation, laccase, lignin peroxidase and Mn-containing peroxidases
were identified. The products of dye metabolism exhibited low toxicity.
A synergistic interaction has been shown between cells of the yeast Rhodotorula sp., in
a consortium with the bacteria Raoultella planticola and Staphylococcus xylosus, in the process
of biodegradation of methylene blue [51]. Intracellular NADH and DCIP reductases were
the key enzymes involved in the process of dye degradation. In addition, azoreductase,
tyrosinase, laccase, nitrate reductase, MnP and LiP also contributed significantly to the
degradation process of methylene blue. The authors of the study confirmed the phytotoxic-
ity and cytotoxicity of dye degradation by-products. When cells of the same consortium
immobilized in a Ca-alginate gel were used to purify municipal wastewater and industrial
effluents contaminated with the same dye (200 mg/L), the degradation efficiency of methy-
lene blue reached 100% and 78.5%, respectively, when treatment was carried out within
144 h.
A mixture of 15 azo dyes (20 mg/L of each type) was subjected to microbial degra-
dation under the influence of fungal (A. niger, A. terrus, A. oryzae and A. fumigatus) and
bacterial (Brevibacillus brevis, Bacillus coagulans, Lysinibacillus macroides, L. fusiformis and
B. subtilis) consortia [52]. The most active bacterial strain was B. subtilis, which showed the
highest degree of biodegradation (from 71.8% to 100%) of eight azo dyes. However, under
the influence of the bacterial consortium, the degradation efficiency was lower than that of
the fungal consortium, which turned out to be capable of biodegradation of all 15 tested
azo dyes. At the same time, 10 azo dyes were degraded by 100% under the action of the
fungal consortium.
The anthraquinone dye Disperse Red 3B (0.1 g/L) was decomposed by a consortium
consisting of cells of the fungus Aspergillus sp. and microalgae Chlorella sorokiniana for
4 days, with an efficiency of 98.1% [53]. In this case, the resulting low-molecular-weight
dye destruction products were characterized by low toxicity. When the dye concentration
increased by five times, the bleaching efficiency decreased to 69.3%, while when consortium
cells were used separately, the bleaching efficiency was almost seven times lower (less
than 10%).
The efficiency of methyl orange bleaching by mixed cultures of the fungus Daedalea
dickinsii and bacteria P. aeruginosa reached 98% within 7 days [54], while the efficiency of
dye degradation by one fungus was only 67.5%.
To degrade azo dyes, a yeast consortium consisting of two cultures (Sterigmatomyces
halophilus and Meyerozyma guilliermondii) was obtained [55]. In optimal conditions (pH
7.0, 35 ◦ C and 50 mg/L dyes), the efficiency of degradation of dyes (Reactive Black 5,
Acid Orange 7; Reactive Green 19, Reactive Yellow, ABC, Atlantic Black C) was 88–97%.
Moreover, NADH--DCIP-reductase and azoreductase were the main enzymes involved in
this process.
A consortium consisting of two filamentous fungi (Penicillium oxalicum and Aspergillus
tubeingensis) provided effective degradation (96%) of Congo red within 12 h, which was sig-
nificantly better when compared to the same individual strains (30–69%) [56]. Presumably,
the increased degradation could be the result of mutualism between the two cultures. The
products obtained as a result of the destruction of Congo red were non-toxic for the growth
of microorganisms and plants. An increase in the degree of dye degradation positively
correlated with the production of laccase and Mn-containing peroxidase by fungi.
Thus, when using microbial consortia containing fungi or yeast, it is possible to
effectively decompose dyes, including those present in the mixture in real wastewater. The
main role (in such bioremediation of water sources contaminated with various dyes) is
played by complexes of oxidoreductases secreted by fungal cells.
2.3. Destruction of Synthetic Polymers

Synthetic polymers, also known as man-made polymers, are polymers that are ar-
tificially produced by humans. Synthetic polymers can be classified into four distinct
categories, namely thermoplastics, thermosets, elastomers and synthetic fibers. Synthetic
polymers are derived from petroleum oil. The most common types of synthetic polymers
are: polyethylene, polypropylene, polyvinyl chloride, polystyrene, polyurethane, polyvinyl
chloride, polyethylene terephthalate [7].
The use and production of polymer materials is constantly increasing, and the amount
of waste that enters the environment is increasing at the same time [57]. During the
process of degradation, the size of plastic particles decreases, increasing the possibility
of them penetrating into the cells of microorganisms, animals and plants, thus creating
their potential toxicity. In addition, heavy metals, polycyclic aromatic hydrocarbons,
pesticides, pharmaceutical compounds and other pollutants can be adsorbed onto plastic
microparticles [58]. Microplastics are tiny plastic particles less than 5 mm in size [7]. It
has been found that microplastic particles can change the formation of soil aggregates
and the bulk density and water-holding capacity of the soil, which ultimately leads to
changes in the composition of microbial communities and a decrease in plant growth and
development [59].
Today, many works study the destruction of microplastics by using physicochemical
(thermoconversion, hydrogenolysis, silylation, electrooxidation and photolysis, solvolysis)
and biocatalytic (enzymatic hydrolysis) methods [7]. However, physicochemical processes
are expensive, and require high temperatures and energy, and need special equipment
that uses strong acid or metal catalysts; in addition, the range of microplastics effectively
decomposed by individual enzymes is limited to polyesters and polyamides [7]. As a result,
microbial degradation of polymers remains a relevant preoccupation for researchers, with
artificial consortia generating maximum interest (Table 3) [60–67].
Table 3. Microbial consortia with fungal species for degradation of synthetic polymers.

Sterigmatomyces halophilus, Low-density polyethylene (LDPE)
30 ◦ C, 45 days
Meyerozyma guilliermondii, M. caribbica [60] mass reduction—33.2%
A. niger, P. aeruginosa [61] 37 ◦ C, 30 days Polyurethane weight loss—20%
Curvularia lunata, Alternaria alternata, Penicillium
90 days Polyethylene weight loss—27%
simplicissimum, Fusarium sp. [62]
A. niger, A. flavus, A. oryzae [63] 55 days Polyethylene weight loss—26.2%
Microorganisms isolated from activated sludge and
river sediments (Lysinibacillus massiliensis, Bacillus 160 rpm, 56 days at room Weight loss of sample (LDPE &
licheniformis, B. indicus, B. megaterium, temperature, thermoplastic starch &
B. cereus, Pseudomonas alcaligenes, Aspergillus sp., 10 mL of bacterial and styrene-ethylene-styrene)—16%
Penicillium sp., Alternaria sp., Candida parapsilosis [64] fungi suspension, and one film
Microorganisms isolated from compost (B. sonorensis, sample (1 cm2 ) of
B. subtilis, Aspergillus sp. Trichoderma sp., polymer materials Weight loss—21.9%
Rhizopus sp.) [64]
Microorganisms of enriched landfill soil
(Achromobacter xylosoxidans, Trichosporon pH 7.2, 150 rpm, 30 ◦ C, 90 days LDPE weight loss—55.6%
chiropterorum, Penicillium chalabudae) [65]
Polypropylene/poly (butylene
29 ◦ C, 85% humidity, adipate-co-
Aspergillus sp., Penicillium sp. [66]
30 days terephthalate)/thermoplastic starch
weight loss—1.0–2.3%
Bacillus sp., Aspergillus sp. [67] 30 ◦ C, 150 rpm, 30 days LDPE weight loss—12%
Biodegradation of microplastics is analyzed, above all, by determining the weight

loss of a polymer sample, changes in its mechanical properties, and/or the appearance
of destruction products. More than 90 microorganisms, including fungi, have been iden-
tified that are capable of degrading synthetic polymers [68,69]. It should be noted that
biodegradation is a long-term process. It is possible to intensify it through pretreatment of
the original polymers that uses chemical or physical methods [70].
More efficient degradation of plastics can only be achieved by the action of microbial
consortia, since complexes of enzymes that cannot be synthesized by a single microorganism
are required. For example, to achieve the degradation of low-density polyethylene (LDPE),
a consortium of three yeast strains Sterigmatomyces halophilus, Meyerozyma guilliermondii
and M. caribbica was obtained [60], which provided a decrease in the strength of the
polymer and weight loss by 63.4% and 33.2%, respectively, within 45 days The yeast were
capable of synthesizing LDPE-degrading enzymes (such as Mn peroxidase, laccase, and
lignin peroxidase), whose activity in the consortium increased. The development of the
biodegradation process was accompanied by the active accumulation of the biomass of
yeast destructor cells, and the products of polymer metabolism were alkanes, alkenes,
carboxylic acids, aldehydes, ethers and alcohols.
The process of biodegradation of polyurethane by a consortium consisting of cells
of the fungus A. niger and bacteria P. aeruginosa was studied [61]. Cellulose (4 wt%) was
added to the medium to stimulate the biodegradation of polyurethane by intensifying the
induced synthesis of the necessary enzymes and, as a result, the weight loss of polymer
was about 20%.
A fungal consortium consisting of strains of Curvularia lunata, Alternaria alternata,
Penicillium simplicissimum and Fusarium sp. degraded polyethylene, producing 27% weight
loss within 3 months. Degradation by individual strains of polyethylene, meanwhile,
was low and amounted to only 0.7–7.7% [62]. Similar results were obtained during the
biodegradation of LDPE by a consortium consisting of three strains of fungi of the genus
Aspergillus (A. niger, A. flavus and A. oryzae), with weight loss of polymer about 26% during
55 days [63].
Consortia of microorganisms obtained by isolating the most active strains from various
sources (activated sludge, compost and river bottom sediments) were used to compile
consortia and degrade polymeric materials [64]. The isolated microorganisms were bacteria
of the genus Bacillus and Pseudomonas, fungi of the genera Aspergillus, Rhizopus, Alternaria,
Penicillium and Trichoderma, as well as the yeast Candida parapsilosis. A polymer sample
consisting of LDPE, thermoplastic starch and styrene-ethylene-styrene was biodegraded by
the resulting consortia, with 16–22% weight loss within 56 days.
Microbial consortia, composed of microorganisms isolated from enriched landfill soil,
were used to degrade LDPE for 90 days, resulting in polymer weight reduction to 55.6% [65].
The authors attribute the reason for such a long process, with a relatively low percentage of
polymer destruction, to the large composition of the consortium.
The effect of extrusion cycles on the degradation of a polypropylene/poly(butylene
adipate-co-terephthalate)/thermoplastic starch mixture by a fungal consortium consisting of
only two filamentous fungi (Aspergillus sp. and Penicillium sp.) was studied for 30 days [66].
Extrusion for seven cycles increased the biodegradation efficiency by two times. However,
the weight loss of the polymer was still insignificant (2.3%).
Individual microorganisms (bacteria Bacillus sp. and fungi Aspergillus sp.) were in-
troduced into a consortium degraded LDPE with an efficiency of 10%, while a microbial
consortium consisting of the same bacteria and fungi showed slightly more efficient degra-
dation of the same polymer as a whole (12%). However, no antagonistic effect was observed
between the cells of the consortium [67].
In general, it is worth noting that the biological degradation of synthetic polymers is
still a long process, despite the constant search for new solutions in this area, including
the use of new microbial consortia. In this regard, it may be more effective to combine
physicochemical and biological methods with the use of microbial and enzymatic catalysts.
2.4. Degradation of Pesticides

Pesticides are widely used in agriculture to control insects, weeds, pathogenic fungi
and bacteria. Excessive and uncontrolled use of pesticides leads to serious environmental
problems, including their accumulation in soil and water sources [71]. In some cases, even
after wastewater treatment at the treatment plants, the concentrations of pesticides can
remain quite high [72]. The use of biological methods is considered to be an effective
approach to the degradation of pesticides, but the efficiency of such processes may be
low and they may have a significant duration [73]. For this reason, the search for new
strategies for the biocatalytic elimination of pesticides continues to be relevant, and the
use of microbial consortia, including those containing fungi, seems to offer a promising
solution to this problem (Table 4 [74–79]).
The use of a consortium of the bacteria Bacillus subtilis and the fungus Fomitopsis pinicola
to achieve the biodegradation of the pesticide DDT (1,1,1-trichloro-2,2-bis(4-chlorophenyl)
ethane) is known. Within 7 days at 30 ◦ C, the degree of pesticide degradation reached
86% [74]. Similar results for the degradation of DDT, when the efficiency of the process was
86%, were obtained using a consortium consisting of the fungus Pleurotus ostreatus and the
bacteria Pseudomonas aeruginosa [75].
Table 4. Microbial consortia with fungal species for degradation of pesticides.

DDT (1,1,1-trichloro-2,2-bis(4-
Fomitopsis pinicola, B. subtilis [74] 30 ◦ C, 7 days chlorophenyl) ethane)
degradation—86%
Pleurotus ostreatus, P. aeruginosa [75] 25 ◦ C, 7 days DDT degradation—86%
38 pesticides in mixture—total
concentration—72.7µg/L,
A. niger, Chlorella vulgaris [76] Degradation—23%
biomass—181.6 mg dry weight/L, pH 4.0,
100 rpm, 68 h
Concentration of each pesticide—50 mg/L, Degradation of atrazine—81%,
Verticilium sp., Metacordyceps sp. [77]
100 rpm, pH 5.5, 27 ◦ C, 21 days iprodione—96%; chlorpyrifos—99%
Concentration of each pesticide—50 mg/L, Degradation of atrazine—64%,
Verticilium sp., Metacordyceps sp.
flow rate—90 mL/h, inoculum iprodione—96%; chlorpyrifos—85%
immobilized in Ca-alginate beads [77]
concentration—30 w/v, 100 rpm, 27 ◦ C (11–15 days)
Consortium of microorganisms present in Degradation of atrazine—72.2%,
coconut fiber, garden compost and Mixture of pesticides—30–40 mg/kg, pH 6.4, carbendazim—96.7%,
agricultural soil and 25 ◦ C, 16 days carbofuran—98.7%,
Trametes versicolor [78] metalaxyl—96.7%
Fomitopsis pinicola, Ralstonia pickettii [79] DDT—5 mM, 30 ◦ C, 7 days DDT degradation—61%
A consortium of cells from the microalgae Chlorella vulgaris and the fungus A. niger was
used to remove pesticides from water. When treating a mixture of 38 pesticides at a total
concentration of 72.7 µg/L, the biodegradation efficiency reached 23%. Moreover, the final
concentrations of some pesticides (difenoconazole, carfentrazone ethyl, phenmedipham
and trinexapac ethyl) were, after treatment by the consortium, even below the detection
limit [76].
The efficiency of biodegradation of pesticides (atrazine, iprodione, chlorpyrifos),
under the action of a suspended fungal consortium consisting of Verticilium sp. cells and
Metacordyceps sp. [77], reached 81–99% within 21 days. Laccase, MnP, and Mn-independent
peroxidase were the main enzymes involved in pesticide degradation. The use of the
same consortium in an immobilized form (incorporation into a Ca-alginate gel) for the
degradation of pesticides, in a packed-bed bioreactor in a continuous mode, ensured that,
at a medium flow rate of 90 mL/h, 64–96% of pesticide degradation was achieved. The
performance of the process depended on many factors, including the flow rate of the
medium, the type of pesticide itself and the time of its processing.
The efficiency of biodegradation of pesticides (atrazine, carbendazim, carbofuran,
metalaxyl) by a microbial consortium (composed of microorganisms present in coconut
fiber, garden compost and agricultural soil) supplemented with the fungus Trametes versi-
color was assessed [78]. This consortium effectively biodegraded all tested pesticides by
72–99% within 16 days. The addition of the antibiotic oxytetracycline (10 mg/kg) did not
significantly affect the efficiency of pesticide degradation by this consortium.
A synergistic effect was observed during the degradation of DDT by a mixed con-
sortium composed of the fungi Fomitopsis pinicola and bacteria Ralstonia pickettii [79]. The
degradation of the pesticide after 7 days was 61%; when the same microorganisms was
used separately; the degradation of DDT was 31–42%. Degradation of DDT occurred
through its transformation into 1,1-dichloro-2,2-bis(4-chlorophenyl)ethane, which was
further transformed into 1,1-dichloro-2,2-bis(4-chlorophenyl)ethylene, and eventually con-
verted to 1-chloro-2,2-bis(4-chlorophenyl) ethylene. These metabolites were less toxic than
DDT. Fungi F. pinicola produced extracellular enzymes, such as oxidoreductase, super-

oxide dismutase, catalase, laccase and cytochrome P450 monooxygenase. Cells of the
bacteria R. pickettii synthesized monooxygenase, lipase, and depolymerase. F. pinicola and
R. pickettii, which were shown to be in co-metabolism, since F. pinicola fungi converted
DDT into products that were used by bacterial cells. When fungal cells grew together with
bacteria, the formation of a thick layer of hyphae was observed, indicating that the bacterial
cells stimulated mycelial growth.
In general, the presence of fungi in mixed artificial consortia makes it possible to more
effectively catalyze the decomposition of pesticides (both individually and in mixtures), in
comparison with individual cultures. However, concerns arise about the possible formation
of resistant strains meaning that, along with the creation of consortia, it is necessary to
develop methods for regulating their composition and metabolic activity.
2.5. Degradation of Polycyclic Aromatic Hydrocarbons

Polycyclic aromatic hydrocarbons (PAH), formed as a result of human anthropogenic
activities, constitute a major class of environmental pollutants, some of which are toxic and
resistant to degradation [80].
Along with PAH, other contaminants (heavy metals, pesticides and other pollutants)
may be simultaneously present, which can lead to an increase in the overall toxicity of the
media and, most importantly, complicate bioremediation. Various physicochemical meth-
ods for the remediation of PAHs are currently available, but they have several disadvan-
tages, including high cost, difficulty in implementation, and, sometimes, inefficiency [81].
Various individual strains of microorganisms, including microalgae, fungi and bacte-
ria, have demonstrated the ability to biodegrade PAHs [82]. Bacterial destruction of PAHs
consists of their enzymatic transformation under the action of monooxygenases or dioxyge-
nases with the formation of intermediate compounds such as catechols. Some fungi of the
genera Cunninghamella, Coriolopsis, Chrysosporium, Trichoderma, Phanerochaete, Aspergillus,
Pleurotus, Trametes, Bjerkandera, Penicillium, Mucor and Cladosporium can metabolize PAHs
as a sole carbon source or co-metabolize them when using other substrates.
Microalgae have various mechanisms for the decomposition or removal of PAHs,
namely biosorption, bioaccumulation, biodegradation, and complexation [83]. The biodegra-
dation of PAHs by ligninolytic and non-ligninolytic fungi is characterized by two different
main mechanisms that occur due to various enzymes mainly involved in these processes.
Secreted laccase, MnP, and lignin peroxidases of ligninolytic fungi are the most active
catalysts acting on the aromatic rings of PAHs [84]. Oxidases can participate in the for-
mation of free hydroxyl radicals by giving up one electron, which oxidizes PAH rings
outside fungal cells. The products of such reactions are PAH-quinones and acids, whereas
intracellular cytochrome P450 monooxygenases, and hydroxylating low molecular weight
PAHs, are the main participants in the oxidative degradation of PAHs carried out by non-
ligninolytic fungi [84]. Cytochrome P450 monooxygenases oxidize PAHs to epoxides and
dihydrodiols. However, in artificial microbial consortia of mixed composition, a synergistic
metabolic effect can develop, leading to improved degradation of PAHs by consolidating
various catalytic mechanisms (Table 5 [85–92]). As has been noted [81], PAHs are often
present in the environment as a mixture, and bioremediation can therefore occur through
co-metabolism. For example, for the biodegradation of PAHs in the presence of heavy
metals, a microbial consortium consisting of B. subtilis and the fungus Acremonium sp.
was used, which effectively decomposed (61–100%) naphthalene, fluorine, phenanthrene,
anthracene and fluoranthene within 10 days [85].
The study of the biodegradation of benzo[a]pyrene by cells of the fungus Pleurotus
ostreatus, as well as consortia of this fungus with Penicillium chrysogenum or with the bacteria
P. aeruginosa is known. The efficiency of benzo[a]pyrene degradation by consortia (86.1 and
75.1%, respectively) was higher than that of an individual fungal strain (64%) [86].
A mixed consortium was created, in which cells were immobilized on biochar to
degrade phenanthrene (50 mg/L) while removing 150 mg/L Cd2+ . The dominant species
were bacteria Proteobacteria and Bacteroidota, and fungi of Fusarium genus [87]. The immobi-
lized consortium effectively decomposed phenanthrene (up to 98%) within 7 days, while
simultaneously removing up to 99% of Cd2+ . Immobilization of the consortium improved
its dehydrogenase activity, which increased the degradation of phenanthrene. In addition,
immobilization led to stabilization of the composition and more favorable development of
the microbial community in the consortium.
Table 5. Microbial consortia with fungal species for degradation of PAHs.

Concentration of each PAH in Degradation of naphthalene—100%,
Acremonium sp, B. subtilis [85] mixture—50 mg/L, 28 ◦ C, 160 rpm, fluorine—89%, phenanthrene—82%,
10 days anthracene—71%, fluoranthene—61%
Pleurotus ostreatus, Penicillium
Degradation of benzo[a]pyrene—86%
chrysogenum [86] 30 ◦ C, 30 days
P. ostreatus, P. aeruginosa [86] Degradation of benzo[a]pyrene—75%
Consortium (Proteobacteria, Bacteroidota, Mixture of 50 mg/L of phenanthrene and Degradation of phenanthrene—92–98%,
Fusarium) immobilized on biochar [87] 150 mg/L of Cd2+ , 150 rpm, 30 ◦ C, 7 days removing of Cd2+ —94–99%
Mixture of pyrene and Degradation efficiency of
Consortium with two genetically
benzo(a)pyrene—1000 mg/kg soil, pH of phenanthrene—92%, pyrene—64%,
modified strains of A. niger [88]
8.4, 30 ◦ C, 14 days benzo(a)pyrene—65%
Mixture of naphthalene, fluorene,
Kocuria rosea and A. sydowii immobilized
phenanthrene, anthracene, and
in guargum-nanobentonite composite Degradation efficiency—85–100%
pyrene—100 µg of each PHA/g of soil,
water dispersible granules [89]
pH 8.3, 27 ◦ C, 30 days
800 mg/L of pyrene, rhamnolipid
P. putida, yeast Basidioascus persicus [90] Degradation efficiency—78%
biosurfactant 100 µL, 28 ◦ C, 21 days
Degradation of fluoranthene,
indene[1,2,3-cd]pyrene and
Concentrations of different
Ochrobactrum intermedium and white rot benzo[g,h,i]perylene—100%;
PAH—138.2–268.0 mg/kg of soil,
fungus Pleurotus ostreatus [91] Anthracene, pyrene, chrysene and
moisture—70%, 30 ◦ C, 110 days
benzo[a]anthracene—96%, 86%, 98% and
98%, respectively
A mixture of anthracene, phenanthrene,
Pleurotus ostreatus, fluorene, pyrene, and
Degradation efficiency > 70%
Azospirillum brasilense [92] fluoranthene—50 mg/L, 130 rpm, 24 ◦ C,
14 days
Using the method of induced microbial selection, microbial consortia were created,
including natural fungal (Aspergillus flavus, A. nomius, Rhizomucor variabilis, Trichoderma
asperellum, and A. fumigatus) and bacterial strains (Klebsiella pneumoniae, Enterobacter sp.,
Bacillus cereus, Pseudomonas aeruginosa, Streptomyces sp., Klebsiella sp., and Stenotrophomonas
maltophilia), as well as genetically engineered strains of A. niger expressing lignin peroxidase
and MnP genes for the degradation of phenanthrene, pyrene and benzo[a]pyrene in soil [88].
The efficiency of degradation of a mixture of PAHs present in the soil at a concentration of
1 g/kg under the influence of this consortium reached 65–92% after 14 days of the process.
Cells of a microbial consortium consisting of the bacteria Kocuria rosea and the fungus
A. sydowii were used after immobilization in guargum-nanobentonite composite water
dispersible granules to degrade a mixture of PAHs (naphthalene, fluorene, phenanthrene,
anthracene, and pyrene) containing 100 µg/g of each pollutant in sandy loam soil. Cell
immobilization ensured PAH degradation by 85–100% [89].
An increase in the efficiency of pyrene degradation (up to 78%) was noted when
using a microbial consortium consisting of bacteria P. putida and yeast B. persicus, when
rhamnolipid was added to the medium as a biosurfactant. The use of rhamnolipid led to
an increase in the bioavailability of pyrene [90].
A microbial consortium including Ochrobactrum intermedium and Pleurotus ostreatus

was able to degrade a mixture of PAHs in soil [91]. The consortium completely removed
fluoranthene, indene[1,2,3-cd]pyrene and benzo[g,h,i]perylene within 50, 80 and 50 days,
respectively. Anthracene, pyrene, chrysene and benzo[a]anthracene were degraded by
86–98% in 110 days. The consortium was more effective than the two microorganisms
used alone. Bacteria formed biofilms around fungal hyphae, stabilizing the high functional
activity of the consortium.
The degradation of PAHs (anthracene, phenanthrene, fluorene, pyrene, and fluoran-
then) was studied using a mixed consortium containing P. ostreatus and A. brasilense [92].
Compared to monocultures (20–60%), the efficiency of PAH degradation by the consortium
exceeded 70%, due to an increase in the synthesis of extracellular laccase and versatile
peroxidase. At the same time, an increase in the mycelium biomass and the number
of bacterial cells was found in the consortium, which confirmed the utilization of PAH
destruction products.
In analyzing the presented data, one can note the high efficiency of mixed consortia in
biocatalytic processes of PAH decomposition. It turned out that it was possible to increase
the efficiency of consortia, including through the use of genetically improved strains [88].
However, despite the efficient functioning of recombinant cells, their use is associated with
a number of problems, with the need to maintain selective cultivation conditions for them
presenting itself as a particular problem. It is also possible to increase the efficiency of
degradation by using immobilized forms of consortia [87,89]. In this case, it is advisable to
use the methods of incorporating cells into gel matrices and sorption of cells on various
carriers as the main methods of immobilization.
2.6. Degradation of Pharmaceutical Pollutants

The active use of various pharmaceutical compounds (antibiotics, hormones, anal-
gesics, anticonvulsants, anti-inflammatory, cardiovascular, antiepileptic and other drugs)
has led to them contaminating environmental objects. The presence of these substances in
wastewater is detected in concentrations from 0.15 ng/L to 2.0 g/L [93]. These compounds
are most often highly toxic and resistant to decomposition, and can therefore accumulate,
causing harm to microorganisms and various animals.
Physicochemical processes (membrane filtration, ozonation, and photocatalytic oxida-
tion, sorption, etc.) do not provide complete removal of pharmaceutically active compounds
(PhACs) and are characterized by a fairly high cost, the formation of toxic by-products, low
selectivity, etc. [6].
The biocatalytic transformation of PhACs under the action of enzymes such as laccase
and peroxidases has become widespread. However, a number of problems limit their use
including, in particular, the low activity of laccases at neutral pH values or the need to add
H2 O2 as a substrate for peroxidases. In addition, it is necessary to ensure the effective and
long-term functioning of enzymes, which is achieved by immobilizing them and increasing
the cost of the resulting enzyme preparations [6].
It has also been established that the efficiency of enzymatic hydrolysis sharply de-
creases when treating real wastewater, which is because the micropollutants and solid
suspended microparticles can significantly reduce the activity of enzymes, acting as their
inhibitors and sorbents [6]. In addition, enzyme complexes are needed. In this regard,
microbiological wastewater treatment is still relevant [94], and the use of artificial consortia
for the bioremediation of PhACs seems to be the most effective approach, as this ensures
the simultaneous synthesis by different cells of microorganisms of different enzymes in-
volved in the destruction of pollutants (Table 6 [95–101]). In particular, in an artificial
consortium of fungi consisting of Phanerochaete chrysosporium and Pycnoporus sanguineus,
100% degradation of ciprofloxacin, norfloxacin and sulfamethoxazole (SMX) was shown,
due to the presence of a complex of oxidative enzymes (laccase, lignin peroxidase and Mn
peroxidase) [95]. When a pure culture of P. chrysosporium was used, the efficiency of antibi-
otic degradation was lower (64.5%, 73.2% and 63.3%, respectively), and the adsorption of
antibiotics by fungal mycelium in the consortium was only 2–3%.
Table 6. Microbial consortia with fungal species for degradation of pharmaceutical pollutants.

Each antibiotic concentration—10 mg/L,
Removal efficiency of ciprofloxacin,
Pycnoporus sanguineus, Phanerochaete biomass of each
norfloxacin and sulfamethoxazole in their
chrysosporium [95] strain—0.15 g dry weight/L), pH 4.5,
mixture—100%
30 ◦ C, 4 days
Sulfamethoxazole (50 mg/L) and
Pycnoporus sanguineus,
vitamins mixture (VB2, VB6, VB12 and Sulfamethoxazole degradation—93%
Alcaligenes faecalis [96]
VC), 28 ◦ C, 120 rpm, 24 h
Concentration of each of three
Ganoderma applanatum, Laetiporus Degradation (mixture of celecoxib,
pollutants—10 mg/L, pH 6.4, ambient
sulphureus [97] diclofenac and ibuprofen)—99.5%
temperature, 150 rpm, 72 h
A. niger, Mucor circinelloides, Trichoderma Pollutants concentration—1 mg/L, pH Degradation of carbamazepine—90%,
longibrachiatum, Trametes polyzona and 4.6, 30 ◦ C, 7 days, consortium diclofenac sodium—96% and
Rhizopus microsporus [98] concentration—30% (v/v) ibuprofen—91%
Pharmaceutical substances—8–11 µg/L,
Relative removal of initial ranitidine
A. niger, C. vulgaris [99] microalgae-fungus
concentrations—50%
biomass—75 mg dry weight/L, 72 h
Penicillium rastrickii, P. oxalicum,
Mixture of diclofenac, carbamazepine
Cladosporium cladosporoides, Micrococcus Degradation of diclofenac—99%,
and ketoprofen with 100 µM of each
yunnanensis, Oligella ureolytica, ketoprofen—80%
compound, 28 ◦ C, 10 days
Sphingobacterium jejuense [100]
Simulated swine wastewater with Removal efficiency of
addition of 0.1–0.5 mg/L Cu (II), sulfamonomethoxine, sulfamethoxazole
Chlorella vulgaris, Aspergillus oryzae [101]
0.4 mg/L of mixture of antimicrobial and sulfamethazine—58.8%, 63.5%, and
agents, pH 7.2, 28 ◦ C, 14 days 63.9%, respectively
SMX has been shown to significantly inhibit the laccase activity of the fungus
P. sanguineus, which reduces the biodegradation efficiency of this antibiotic. To elimi-
nate this problem, a consortium of this fungus with bacterial cells of Alcaligenes faecalis
was used [96]. The removal efficiency of SMX within 24 h by the consortium in a medium
containing a mixture of vitamins (VB2, VB6, VB12, and VC) was higher (93%) than when
only pure fungal culture was used (28%). It was confirmed that the fungal mycelium
adsorbs only 1% SMX—that is, the reduction in antibiotic concentration was achieved
precisely due to biotransformation.
Biodegradation of a mixture of nonsteroidal anti-inflammatory drugs (celecoxib, di-
clofenac and ibuprofen) was carried out under the influence of two fungal strains Ganoderma
applanatum and Laetiporus sulfureus for 72 h [97]. The efficiency of pollutant removal by the
consortium was 99.5%, while the same microorganisms individually provided 61 and 73%
degradation, respectively. A significantly increased induction of synthesis of a number of
enzymes (laccase, lignin peroxidase and MnP) in the consortium was revealed (by 201, 180
and 135%, respectively).
Degradation of carbamazepine, diclofenac and ibuprofen (1 mg/L) was carried out by a
multicomponent fungal consortium (A. niger, Mucor circinelloides, Trichoderma longibrachiatum,
Trametes polyzona and Rhizopus microsporus) in a sequencing batch reactor [98]. It was shown
that the enzymatic activity of laccase, MnP and lignin peroxidase increases with increased
biomass. The efficiency of pollutant removal within 7 days was more than 90% (Table 6).
Water purification from seven pharmaceutical drugs (acetaminophen, carbamazepine,
diclofenac, metoprolol, naproxen, ranitidine and SMX) was carried out using cells of
microalgae C. vulgaris, the fungus A. niger and their consortium. After 72 h, and only when
the consortium was used, a decrease in the initial concentration of ranitidine by 50% was
noted. For other substances, the purification efficiency was insignificant, which was due to
the low concentration of the used consortium in the reaction medium (it was 16–500 times
lower than what is usually used in wastewater treatment plants) [99].
Various microorganisms (fungi and bacteria) isolated from sewage sludge were studied
to obtain artificial microbial consortia capable of effectively degrading pharmaceutical
pollutants (diclofenac, carbamazepine and ketoprofen). It was shown that consortia with
the fungi Penicillium oxalicum and Penicillium rastrickii are able to effectively degrade
diclofenac (99%) and ketoprofen (80%) within 10 days. At the same time, under the
influence of individual crops, the efficiency of degradation was low [100]. It is important to
note here that while the consortia showed high efficiency, their ability to degrade pollutants
could be suppressed by the appearance of “negatively” acting species of microorganisms,
leading to the cessation of interactions between the components of the consortia and their
trophic connections. Thus, such a “negative” action of microorganisms on the consortium
at a certain stage can be considered as a regulator of its bioremediation activity that allows
it to slow down the speed of functioning of such artificial cellular biosystems, if necessary.
A consortium based on the microalgae C. vulgaris and the fungi A. oryzae was used for
the biodegradation of a mixture of antibiotics (sulfamonomethoxine, SMX and
sulfamethazine) [101]. At the same time, different concentrations of copper (Cu(II)) were in-
troduced into the medium. The results confirmed that the addition of Cu(II) (0.1–0.5 mg/L)
increased the removal of sulfamonomethoxine (58.8%), SMX (63.5%) and sulfamethazine
(63.9%). The degradation pathways of these compounds were associated with hydroxy-
lation, deamination, oxidation, desulfurization, and the breaking of chemical bonds. The
fungal mycelium sorptionally retained microalgae cells on its surface, helping to stabilize
the consortium. Copper ions had a great influence on the accumulation of microalgae
biomass, since they can absorb and use Cu(II) in photosynthesis (instead of magnesium) in
the chlorophyll structure, continuing to fix CO2 and synthesize carbohydrates. In addition,
it was found that Cu(II) increased the lipid content of Chlorella cells in the co-culture system.
These examples indicate the high efficiency (50–100%) of the use of artificial microbial
consortia with fungal cells in the processes of bioremediation of wastewater contaminated
with pharmaceutical pollutants. At the same time, mainly fungal oxidoreductases are
involved in the implementation of the decomposition mechanisms of pollutants.
2.7. Elimination of Pollutant Mixtures

Today, pollution of various environmental objects with various chemical substances
(natural and synthetic) can have a negative impact on living objects, which is of particular
concern as it raises questions about the need for their analytical control and subsequent
elimination. Here, one of the most effective solutions to the problem seems to be the
use of mixed artificial consortia that include cells of different fungi in their composition
(Table 7 [102–111]).
To clean the soil contaminated with crude oil, a consortium consisting of cells of
the bacteria Acinetobacter baumannii was combined with cells of the fungi Talaromyces
sp. andused. The degree of decomposition of oil was 65.6% after soil treatment for
28 days [102]. In another similar work, not one, but two, bacterial strains (Paraburkholderia
sp. and Paraburkholderia tropica) were combined with the fungi Scedosporium boydii into an
artificial consortium, which was also used for biodegradation of crude oil pollution [103].
Within 1 week, the efficiency of biodegradation reached 81.4% with a mixed consortium.
This parameter was 20% higher, compared to the combined use of two types of bacterial
cells. At the same time, the formation of bacterial biofilms on the surface of the hyphae of
the fungus was revealed, which stabilized the mixed consortium.
Cells of the fungi Scedosporium sp. were combined with bacteria Acinetobacter sp.,
producing biosurfactants for the decomposition of crude oil. The results showed that
the efficiency of oil degradation (200 mg/L) increased from typical for individual mi-
croorganisms, increasing 23–29% to 58.6% within 7 days of an artificial consortium being
used [104]. The ability of bacteria to produce biosurfactants leads to an increase in the
effective biodegradation of hydrophobic substrates.
Table 7. Microbial consortia with fungal species for degradation of various pollutants not mentioned
in Tables 1–6.
Consortia [Reference] * Conditions Pollutant/Process Efficiency

Acinetobacter baumannii, Talaromyces sp. The initial concentration of petroleum in
Degradation of petroleum—65.6%
[102] soil—1220 mg/kg, pH 8.3, 30 ◦ C, 28 days
Paraburkholderia sp., Paraburkholderia
1% v/v crude oil, 120 rpm, 30 ◦ C, 7 days Degradation of crude oil—81.5%
tropica, Scedosporium boydii [103]
Crude oil—200 mg/L, pH 7.0, 150 rpm,
Scedosporium sp., Acinetobacter sp. [104] Crude oil degradation—58.6%
30 ◦ C, 7 days
Greywater—COD—1165.6 mg/L, oil and Degradation of COD, oil and grease and
Micrococcus luteus, Rhodococcus equi,
grease—58 mg/L, sulphate—95.6 mg/L, sulphate were 78.7, 82.6 and
A. niger [105]
pH 7, 35 ◦ C, 96 h 89.7%, respectively
Aspergillus versicolor and bacterial species Greywater with 100 µg/L of carbendazim Degradation of carbendazim and
(Pseudomonas, Klebsiella species, and thiamethoxam, thiamethoxam 94.4 and
B. subtilis) [106] 80 rpm, 30 ◦ C, 240 h 93.6%, respectively
Real textile effluent pH 8.7,
Degradation—78.1%,
A. flavus, Fusarium oxysporium [107] COD—611 mg/L, pH 6.0–8.0, 28 ◦ C,
COD removal—77.6%
7 days
Consortium of Brevibacillus laterosporus
and Galactomyces geotrichum immobilized Textile industry effluent pH 8.8, Degradation during 5 repeated
into Ca-alginate or polyvinyl COD—2400 mg/L, 48–60 h cycles—76–95%
alcohol-alginate beads [108]
Chlorobenzene—220 mg/L, 160 rpm,
Ralstonia pickettii, Trichoderma viride [109] Chlorobenzene degradation—100%
28 ◦ C, 60 h
Poly(vinyl acetate) processing COD, poly(vinyl acetate) and color
Chaetomium globosum, A. niger,
wastewater pH 7.1, COD—23.48 g/L; pH removal yields—97.8%, 98.5% and
Rhizopus oryzae [110]
5.5, 150 rpm, 28 ◦ C, 10 days 99.8%, respectively.
Phenol (1000 mg/L) and selenite Phenol degradation—97.8% with the
Phanerochaete chrysosporium,
concentration—10 mg/L, 180 rpm, pH simultaneous reduction of selenite
Delftia lacustris [111]
6.5, 30 ◦ C, 120 h to Se(0)
* COD—chemical oxygen demand.
Greywater refers to domestic wastewater generated after sewage and washing machine
effluents, which contain various pollutants, including detergents, dyes, heavy metals and
personal care products. A bacterial-fungal consortium with Micrococcus luteus, Rhodococcus
equi and A. niger was used to purify greywater [105]. In optimal operating conditions
(pH 7, 35 ◦ C), the decrease in the content of pollutants (COD, oils, fats and sulfates) was
78.7–89.7% after 96 h of microbial treatment.
In another study, bioremediation of Greywater contaminated with pesticides (car-
bendazim and thiamethoxam) and COD, was carried out under the action of Aspergillus
versicolor fungal cells, a bacterial consortium obtained from kitchen waste sludge, and a
bacterial consortium with added fungi [106]. The efficiency of degradation of COD and
pesticides by the combined consortium was 93.6–94.4% for 10 days, whereas the same
parameter was 44–54% and 84.8–87.9%, respectively, for the bacterial consortium and the
individually applied fungal cells. Thus, the advantages of combining bacterial and fungi
were obvious.
To clean real textile effluents, a fungal consortium consisting of two microorganisms
(Aspergillus flavus and Fusarium oxysporium) was used. After 7 days, the effluent degradation
efficiency was 78%, while the same parameter for individual crops was 52.4–54.7%. The
effectiveness of wastewater treatment was not only assessed by removing dyes, but also by
reducing COD, which amounted to 77.6% [107].
In another study, the effectiveness of degradation of textile effluents was evaluated by
a bacterial yeast consortium (Brevibacillus laterosporus and Galactomyces geotrichum), whose
cells were immobilized in a Ca-alginate gel or a mixed gel of alginate and poly(vinyl
alcohol) [108].
Immobilized cells were able to degrade wastewater for five cycles with an efficiency
of 74–95%. Cells of the same consortium, immobilized on a stainless-steel carrier, provided
90% of wastewater degradation in a reactor with continuous operation at a flow rate of
10 mL/h for three operating cycles [109].
A mixed consortium, including cells of the bacteria Ralstonia pickettii and the fungi
Trichoderma viride, was used in the biodegradation of chlorobenzene. At an initial concen-
tration of 220 mg/L, this compound was completely removed from the medium for 60 h.
The degree of chlorobenzene mineralization, estimated by CO2 yield, was 86.3%. The same
100% removal of the pollutant was much slower when the fungi was used alone (96 h) or
the bacterial cells (72 h). It is interesting to note that this consortium was able to utilize
1000 mg/L chlorobenzene completely during 105 h. Intermediates of chlorobenzene degra-
dation, which inhibit bacterial growth, were easily metabolized by fungal cells, allowing
bacteria to continuously participate in the joint biodegradation process [106].
The bioprocessing of poly(vinyl acetate) containing wastewater was carried out under
the action of cells of a fungal consortium (Chaetomium globosum, Aspergillus niger and
Rhizopus oryzae) [110]. The degree of COD removal and degradation was 97.8% and 99.8%,
respectively. Enzymatic analysis showed that extracellular laccase and lignin peroxidase
participated in biodegradation. In addition, the analysis of metabolic products showed
their mineralization with the formation of formic acid and ethanol.
Simultaneous removal of phenol (1200 mg/L) and selenite (10 mg/L) from synthetic
wastewater was investigated using a consortium composed of Phanerochaete chrysosporium
fungus cells and Delftia lacustris bacteria [111]. It was found that under the action of such a
consortium, selenite ions are biologically reduced to Se(0) nanoparticles with simultaneous
degradation of phenol. In addition, it was found that, over time, bacteria grow and form
a biofilm on the surface of the fungal cells, which does not lead to an improvement, but
rather to a decrease in the efficiency of phenol decomposition. The consortium was already
completely inhibited at a phenol concentration of 200 mg/L.
Thus, the bacterial biofilm limited the mass transfer and respiratory activity of the
fungal culture, generally worsening the functioning of the consortium over time. Therefore,
it is necessary to note here the revealed fact that the characteristics of the consortium
depend on the duration of its operation.
It is obvious that directed association of microorganisms based on the preliminary
selection of active pollutant destructors is an effective approach to the creation of artificial
consortia. When compared to pure cultures, consortia have shown better biodegradation
of specific pollutants or their mixtures that have a complex composition. The presence of
fungal strains in consortia makes it possible to degrade a wider range of pollutants, which
is due to the secretion of enzymes reacting with pollutions. In addition, it is possible to
increase the efficiency of the process by using consortia in an immobilized form.
Immobilized consortia are attractive for use in multiple work cycles, and consortia
of fungi with microalgae have a high potential in the use of pharmaceutical agents in
wastewater treatment [93]. This approach to solving the problems of pollutant removal and
the bioremediation of media is, in comparison to methods of joint cultivation of microalgae
and bacteria, still relatively insufficiently studied.
3. Analysis of Current Trends in the Development of Fungal-Containing Consortia

3.1. Genetically Modified Microorganisms in Artificial Consortia with Fungi
Despite the high efficiency of degradation of various pollutants by natural microbial
consortia [112], there has recently been a growing demand for strains that are improved by
3. Analysis of Current Trends in the Development of Fungal-Containing Consortia
3.1. Genetically Modified Microorganisms in Artificial Consortia with Fungi
Microorganisms 2024, 12, 470

Despite the high efficiency of degradation of various pollutants by natural microbial
17 of 26
consortia [112], there has recently been a growing demand for strains that are improved
by using advanced methods of synthetic biology and metabolic engineering, and
numerical methods in the field of genetic engineering [113]. To date, the use of genetically
using advanced methods of synthetic biology and metabolic engineering, and numerical
modified strains in mixed microbial consortia for the decomposition of various pollutants
methods in the field of genetic engineering [113]. To date, the use of genetically modified
is recognized (Figure 1). The percentage of such consortia is comparable to variants
strains in mixed microbial consortia for the decomposition of various pollutants is recog-
composed of yeast and bacterial cells (Figure 1). One example of the use of such an
nized (Figure 1). The percentage of such consortia is comparable to variants composed of
artificial consortium, as described in Tables 1–7, is the heterologous expression of genes
yeast and bacterial cells (Figure 1). One example of the use of such an artificial consortium,
encoding
as describedMnP and LiP
in Tables 1–7,enzymes in non-ligninolytic
is the heterologous fungi,
expression which
of genes made itMnP
encoding possible
and LiP to
complement the pathway of degradation of PAHs [88], without leaving
enzymes in non-ligninolytic fungi, which made it possible to complement the pathway of toxic
intermediatesofinPAHs
degradation the treated medium.
[88], without leaving toxic intermediates in the treated medium.
Figure 1. The percentage of artificial consortia of different

different composition of the total number (56) of
Tables 1–7.
reviewed studies, presented in Tables 1–7.
However, thethe traditional

traditionalmethod
methodofofrandom
randommutagenesis
mutagenesis is aistime-consuming ap-
a time-consuming
proach,
approach,and searching
and searchingforfor
key
keyamino
aminoacid
acidmutations
mutationsisisextremely
extremelycomplex
complex and
and requires
operating across a large space. With the help of computer modeling, it is possible to
“calculate” enzymes with improved binding affinity and greater specificity of action in
relation to the substrates destruction, which are both necessary [114]; it is also necessary to
successfully introduce producers of “improved” enzymes into consortia, which is another
serious task that confronts researchers.
Another difficulty is that genetically modified strains often have to survive high
concentrations of pollutants [113,115]. Several strategies have been used to improve cellular
tolerance, including changing the composition of membrane lipids, phenotypic screening
through adaptive laboratory evolution, modification of global gene expression, genome
shuffling, directional evolution, and others.
Although genetically modified strains are effective in bioremediation processes [11],
their use is limited to laboratory studies due to minimizing the risks of environmental
impact. Since there are no globally accepted regulatory documents that concern the spread
of genetically modified organisms, the regulation of the development and release of ge-
netically modified organisms varies in different countries, depending on the purposes of
their use, extending from a complete ban on their import, release or use to allowing their
use, subject to varying degrees of regulation. However, despite this, methods of genetically
engineering filamentous fungi continue to be actively developed and used in research
around the world, making it possible to overcome many of the shortcomings of classical
methods for improving strains [115–117]. To eliminate the negative effect of such cells, it
is possible to use several genetic tools, including cell self-destruction systems [118]. Such
approaches to realization of programmed cell death in a certain period of their functioning
can be used and activated after the completion of bioremediation or after accumulation of
certain concentrations of the cells.
3.2. Role of Composition in Artificial Consortia with Fungal Cells

Microbial consortia have a high self-organization, which allows them to carry out the
catalytic conversion of substrates with high efficiency, providing an intensive exchange of
metabolites. Moreover, in consortia, cells have higher adaptability and viability in relation
to environmental factors (pH, temperature, concentration of pollutants, etc.) [119]. Despite
this, most artificial microbial consortia face problems of non-sustainable functioning [120].
Even minor fluctuations in the composition and activities of consortia can have a negative
impact on the effectiveness of the processes taking place with their participation. The
Microorganisms 2024, 12, x FOR PEER REVIEW
difficulty of managing the bioremediation characteristics of artificial consortia is 19 of 28
actually
determined by the qualitative (Figure 1) and quantitative composition (Figure 2) of the
participants in the artificial biosystems being formed.
Theaverage
Figure2.2.The
Figure averageefficiency
efficiencyofofdegradation
degradationofofpollutants
pollutantsby byconsortia
consortiawith
withtwo,
two,three,
three,four
fourandand
more strains in the composition. The percentage of the total number of all options presented inin
more strains in the composition. The percentage of the total number of all options presented
Tables
Tables1–7
1–7(56
(56studies
studiesreviewed)
reviewed)waswascalculated.
calculated.Each
Eachpoint
pointcorresponds
correspondstotothe
theresearch
researchresults
resultslisted
listed
ininTables
Tables1–7.
1–7.
AsAsthe the analysis

analysis of thethedata
datain inTables 1–71–7
Tables showsshowsthat,that,
despite the diversity
despite of the of
the diversity consor-
the
tia being developed
consortia being developed(Figure 1), the ones
(Figure 1), most widely
the ones mostusedwidely
to create effective
used bioremediation
to create effective
microbial systems,
bioremediation that have
microbial demonstrated
systems, that havetheir advantages
demonstrated over
their natural systems,
advantages consist
over natural
of two orconsist
systems, three microorganisms, of which one is aoffungal
of two or three microorganisms, whichculture
one is (Figure
a fungal2).culture
The maximum
(Figure
partThe
2). of all artificial consortia
maximum part of all currently being
artificial developed
consortia combines
currently beingmycelial fungicombines
developed with each
other and with cells of different bacteria (Figure 1).
mycelial fungi with each other and with cells of different bacteria (Figure 1).
ItItisisknown
knownthat thatfungi
fungiand
andbacteria
bacteriaarearecharacterized
characterizedby bydifferent
differentrates
ratesofofsynthesis
synthesisofof
enzymes that are necessary for the catalysis of different processes
enzymes that are necessary for the catalysis of different processes in the bioremediation in the bioremediation
ofofpollutants.
pollutants.Consequently,
Consequently,the therates
ratesofofdifferent
differentprocesses
processesmay maybe becompared
comparedby byvarying
varying
the biomass of certain cells introduced into heterogeneous consortia
the biomass of certain cells introduced into heterogeneous consortia in order to ensure in order to ensure
their maximum effectiveness of action. This is the basis for
their maximum effectiveness of action. This is the basis for the unification of certainthe unification of certain
microorganismsinto
microorganisms intoartificial
artificialconsortia
consortia(Figure
(Figure3).
3).
After analyzing the data in Tables 1–7, it is possible to conclude that there are consortia
consisting of two–three strains of various fungi, mainly mycelial types, that are most
effective in removing 70–90% of heavy metals in mixtures from various media.
For the removal of dyes, the most effective are the consortia consisting of yeast cells,
which provide bioremediation efficiency of up to 100%; the destruction of pesticides is
systems, consist of two or three microorganisms, of which one is a fungal culture (Figure
2). The maximum part of all artificial consortia currently being developed combines
mycelial fungi with each other and with cells of different bacteria (Figure 1).
It is known that fungi and bacteria are characterized by different rates of synthesis of
Microorganisms 2024, 12, 470 enzymes that are necessary for the catalysis of different processes in the bioremediation 19 of 26
of pollutants. Consequently, the rates of different processes may be compared by varying
the biomass of certain cells introduced into heterogeneous consortia in order to ensure
their maximum
the most effectiveness
successful under theofaction
action.
of This is theconsisting
consortia basis for of
thefungal
unification of such
cultures, certain
as
microorganisms into artificial consortia
Verticilium sp. and Metacordyceps sp. (Figure 3).
Figure 3. The average efficiency of the consortia for elimination of different types of pollutants,
calculated on the basis of sources performed in Tables 1–7. Each point corresponds to the research
results listed in Tables 1–7. The six differently colored squares correspond to the six pollutants shown
on the abscissa axis.
Synthetic polymers undergo the most difficult microbial degradation, but the max-
imum weight loss of synthetic polymers (microplastics) is achieved under the action of
consortia that combine bacteria, mycelial fungi or yeast.
In the case of the presence of mixtures of several PAHs in media subjected to biore-
mediation, the most effective (in terms of the degree of degradation of the pollutant
(86–100%)) were consortia that included bacteria and white rot fungus Pleurotus ostreatus,
as well as an immobilized consortium consisting of the bacteria Kocuria rosea and the fungi
Aspergillus sydowii.
Among the most effective in the removal of pharmaceutical pollutants were selected
consortia from cells of white rot saprobic fungus Pycnoporus sanguineus and Phanerochaete
chrysosporium (the removal efficiency of ciprofloxacin-, norfloxacin and sulfamethoxazole
in their mixture was 100%), as well as tinder fungi Ganoderma applanatum and Laetiporus
sulphureus (efficiency of degradation of a mixture containing celecoxib, diclofenac and
ibuprofen was 99.5%).
Bacterial and fungal consortia proved to be the most effective for the treatment of real
wastewater from industrial enterprises and oil pollution. The fungal consortium consisting
of Phanerochaete chrysosporium and Delftia lacustris can be used to remove phenol, with an
efficiency of over 90%.
Of course, in order to achieve the effective functioning of synthetic microbial consortia,
it is important to predict the possible types of interactions of all microorganisms involved
in the functioning of artificially created biosystems [35,36].
Most often, these interactions are based on mutualism or competition [13,121]. Mu-
tualism suggests that jointly cultivated microorganisms have a beneficial effect on each
other, while the composition of artificial consortia is stabilized, but only at a certain cell
density [34,111].
In case of competition for a substrate, consortium members can release metabolites
into the environment that negatively affect other consortium members (organic acids,
mycotoxins, antibiotics, antimicrobial peptides, enzymes) [122–126]. As a rule, these
processes are regulated by cell Quorum Sensing (QS) [16]. QS molecules and the conditions
of their formation can be used to control or regulate the expression of certain genes,
control the composition of consortia, and ensure intercellular connections between certain
consortium members [34,120,127]. In considering this, modern studies of artificial consortia
should enable the creation of model systems for the accumulation of toxic metabolites of
a number of microorganisms (fungi, microalgae, bacteria) and develop effective ways to
detoxify them.
Cell immobilization can also solve the instability problems of microbial consortia,
since cellular communication can be more active in a confined space and positively af-
fects the speed of bioremediation processes. At the same time, the inclusion of cells in
gel matrices [52,77,108] simulates the development of a stabilized state of cells and the
additional formation of biofilms [11,34,91,103,111].
Immobilized fungal consortia can be used in non-sterile conditions at a separate stage,
and then be integrated into conventional waste treatment systems before the action of
aerobic sludge at water processing stations [128–130]. In some cases, the self-stabilization of
the artificial fungi-containing consortia is sufficient for their use in non-immobilized form in
laboratory conditions to treat various real contaminated industrial and environmental water
and soil samples [39,44,49,51,94,107,108,110]. While such studies bring the introduction of
fungal consortia closer to practice, there are not many known examples of pilot or industrial
tests of artificial consortia [131–133]. Some of them follow:
- microbial consortium containing T. versicolor, P. ostreatus, Phanerochaete sp.,
Pseudomonas fluorescens and B. subtilis cells was applied for the treatment of non-
domestic wastewater. This fungal/bacterial consortium was prepared by mixing
fungal biomass pellets with suspensions of bacterial cells. The removal of colored sub-
stances (2700 Color Units550nm ), COD (1.75 g/L) and nitrate (3 mg/L) was 91 ± 2%,
90 ± 4% and 17 ± 2%, respectively, after 15 days of water treatment at a pilot
plant [131];
- consortium of A. niger, Mucor hiemalis and Galactomyces geotrichum, has been tested
for the treatment of real wastewater from industry at a pilot scale station (110 L) and
industrial wastewater treatment plant (1000 L). The efficiency of COD removal in the
industrial reactor was 50% under the influence of this consortium [132];
- consortium containing Acinetobacter oleivorans, Corynebacterium sp., Pseudomonas sp,
Rhodococcus sp., Micrococcus sp. and yeast Yarrowia sp. was tested by Ecophile Co., Ltd.
(Korea) in the biodegradation of hydrocarbons in soil (2300 mg/kg) contaminated with
diesel fuel. This large-scale experiment involved two samples of 100 metric tons of
contaminated soil, both without (control) and with consortium treatment (109 cells/kg
of soil). The introduction of consortium reduced pollution by 57.7% within 2 weeks,
whereas in the control (without the consortium), degradation was only 10.1% [133].
Thus, such positive samples of scaling use of artificial fungal consortia not only
demonstrate their real-world efficacy, but also addresses potential solutions encountered
during practical applications.
4. Conclusions
Today the question of the action efficiency of artificial fungi-containing consortia in the
bioremediation of various pollutants in real conditions remains open and continues to be
studied, despite the accumulating information in proven positive laboratory studies of their
functioning. The stabilization of the complex composition and functional characteristics of
created multicomponent biosystems is one of the key points for the successful scientific
and practical solution of problems in the field of creating new artificial consortia, including
those combining fungi with similar and other types of microorganisms. The analysis of
the accumulated results, their generalization and the identification of common positive
approaches that combine certain microorganisms and organize their use can form the
basis for the development of new processes for the removal of various pollutants that,
with the highest possible level of efficiency, bioremediate various media. The economic
viability and attractiveness of using artificial consortia containing fungi for bioremediation
determines the cost-effectiveness of cultivating individual strains and their availability for
commercial application. The availability can be satisfied by searching for and isolating the
necessary fungi from natural sources or by creating genetically modified strains [116,117].
Whereas the costs of obtaining the necessary amounts of biomass of fungal cells are,
according to the economic estimates just conducted, considered low enough [116,134].
Enzymes synthesized by fungi, the main catalysts of many redox and hydrolytic processes,
are already commercially available, although they are more expensive products, when
compared to the fungal cells producing them. This should be noted when estimating the
economic viability and scalability of using artificial fungi-containing consortia in large-scale
bioremediation projects. The positive aspects of the economy associated with the use of
fungi in various processes, including bioremediation [135], may contribute to the wider
use of artificial consortia with fungi in the near future.
Author Contributions: Conceptualization, E.E.; investigation, E.E., N.S., O.S., A.A., O.M. and
I.L.; data curation E.E. and N.S.; writing—original draft preparation, E.E., N.S., O.S. and A.A.;
writing—review and editing, E.E. and N.S.; supervision, E.E. All authors have read and agreed to the
published version of the manuscript.
Funding: The research was funded by State Task of Lomonosov Moscow State University (121041500039-8).
Conflicts of Interest: The authors declare no conflicts of interest.
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microorganisms

Citation: Efremenko, E.; Stepanov, N.;

Microorganisms 2024, 12, 470. https://doi.org/10.3390/microorganisms12030470 https://www.mdpi.com/journal/microorganisms

- easy-to-use reproducibility of the compositions of the consortia;

2. Main Targets for Bioremediation Based on Various Consortia Containing Fungi

2.1. Removal of Heavy Metals

Consortia [Reference] Conditions Pollutant/Process Efficiency

Bioremediation of heavy metals by microorganisms includes the following main mech-

2.2. Decolorization of Dyes

Table 2. Microbial consortia with fungal species for decolorization of dyes.

Consortia [Reference] Conditions Pollutant/Process Efficiency

2.3. Destruction of Synthetic Polymers

Consortia [Reference] Conditions Pollutant/Process Efficiency

Biodegradation of microplastics is analyzed, above all, by determining the weight

2.4. Degradation of Pesticides

Table 4. Microbial consortia with fungal species for degradation of pesticides.

Consortia [Reference] Conditions Pollutant/Process Efficiency

DDT. Fungi F. pinicola produced extracellular enzymes, such as oxidoreductase, super-

2.5. Degradation of Polycyclic Aromatic Hydrocarbons

Table 5. Microbial consortia with fungal species for degradation of PAHs.

Consortia [Reference] Conditions Pollutant/Process Efficiency

A microbial consortium including Ochrobactrum intermedium and Pleurotus ostreatus

2.6. Degradation of Pharmaceutical Pollutants

Consortia [Reference] Conditions Pollutant/Process Efficiency

2.7. Elimination of Pollutant Mixtures

Consortia [Reference] * Conditions Pollutant/Process Efficiency

3. Analysis of Current Trends in the Development of Fungal-Containing Consortia

Microorganisms 2024, 12, 470

Figure 1. The percentage of artificial consortia of different

However, thethe traditional

3.2. Role of Composition in Artificial Consortia with Fungal Cells

AsAsthe the analysis

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