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Role of Fungi in Bioremediation and Environmental Sustainability

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DOI: 10.1007/978-3-030-54422-5_8

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 Chapter 8 1

Role of Fungi in Bioremediation 2

AU1 and Environmental Sustainability 3

Ajay Tomer, Ramji Singh, Saurabh Kumar Singh, S. A. Dwivedi, 4

Chilkuri Udaykiran Reddy, Malavika Ram Amanthra Keloth, 5
and Riya Rachel 6

Contents 7

8.1 Introduction 000 8

8.2 Mechanisms of Bioremediation 000 9
8.3 Bioremediation of Contaminated Land 000 10
8.4 Bioremediation Potential of Fungi 000 11
8.5 Use of Fungal Enzymes in Bioremediation 000 12
8.6 Mycoremediation Using Fungi 000 13
8.7 Advanced Technologies Used in Fungal Bioremediation 000 14
8.8 Bioremediation Using Fungal Cytochromes 000 15
8.8.1 Factors Affecting Bioremediation 000 16
8.9 Phytoremediation Using Arbuscular Mycorrhizal Fungi 000 17
8.9.1 Fungi as Symbionts 000 18
8.10 Conclusion 000 19
References 000 20

A. Tomer (*) · C. U. Reddy · M. R. A. Keloth · R. Rachel

Department of Plant Pathology, School of Agriculture, Lovely Professional University,
Phagwara, Punjab, India
R. Singh
Department of Plant Pathology, Sardar Vallabhbhi Patel University of Agriculture &
Technology, Meerut, Uttar Pradesh, India
S. K. Singh
Department of Horticulture, Janta Vedic College, Baraut, Uttar Pradesh, India
S. A. Dwivedi
Department of Entomology, School of Agriculture, Lovely Professional University,
Phagwara, Punjab, India

© The Editor(s) (if applicable) and The Author(s), under exclusive license to
Springer Nature Switzerland AG 2021
R. Prasad et al. (eds.), Mycoremediation and Environmental Sustainability,
Fungal Biology, https://doi.org/10.1007/978-3-030-54422-5_8
 A. Tomer et al.

21 8.1 Introduction AU4

22 Bioremediation is an environmentally friendly process using many different

23 microbes to weaken and detoxify harmful pollutants in a parallel or sequential man-
24 ner. Microorganisms (e.g., fungi and bacteria), green plants, or combinations of
25 them used together can convert toxic pollutants into carbon dioxide (CO2) and water
26 (H2O), inorganic salts, microbial biomass, and other products that are less toxic—in
27 effect, accelerating natural metabolic processes that result in these outcomes
28 (Egamberdieva et al. 2008; Gupta and Sinha 2007; Pawar 2012; Prasad et al. 2017, AU5
29 2018; Mohammadi-Sichani et al. 2019).
30 In recent years, interest in exploring microbial biodegradation of toxins has been
31 amplified by human attempts to achieve a sustainable approach to purification and
32 restoration of polluted habitats. Cleaning up polluted soil and water by use of organ-
33 isms—including fungi, bacteria, and their enzymes—is a cost-efficient, sustainable,
34 and natural approach (in comparison with other typical techniques) (Kumar and
35 Dwivedi 2019). In bioremedial technologies, microbes are introduced to improve AU6
36 decomposition or elimination of organic and inorganic pollutants and harmful con-
37 taminants. Pollutant bioremediation can be achieved by various methods such as
38 natural attenuation, biostimulation, bioaugmentation, or combinations of these
39 methods (Bisht et al. 2019). Because of their consistent morphology and versatile
40 metabolic ability, fungi play crucial roles as degraders and symbionts in the envi-
41 ronment as a whole, including soil and aquatic habitats; thus, they are particularly
42 suitable for bioremediation. Mycoremediation is a method of bioremediation using
43 fungi to decontaminate contaminated areas. Arbuscular mycorrhizal fungi (AMFs)
44 primarily perform their functions in soil, achieving and altering the soil microbial
45 balance. AMFs primarily enhance soil microbe growth and restrict plant pathogen
46 proliferation. Because of their symbiotic nature, AMFs depend on plant roots to
47 provide them with the carbon and sugar they need in order to grow and proliferate.
48 Eventually, as the plants and fungi grow together, they both benefit from their asso-
49 ciation. Mycorrhizal combinations make plants soil tolerant, enhance their growth,
50 and keep them healthier than nonmycorrhizal plants. The main division of the book AU7
51 provides an overview of bioremediation and main emphasis of this book is on
52 microbial process because the cycling of organic compounds in the environment is
53 an important part of bioremediation (Zhang et al. 2019).

54 8.2 Mechanisms of Bioremediation

55 Bioremediation is a biological degradation mechanism using microbial capacities to

56 minimize the concentrations and toxicity of a large variety of contaminants, whereby
57 areas contaminated with harmful pollutants are treated with the help of microbial
58 processes. Microorganisms interact physically and chemically with pollutants, lead-
59 ing to structural changes or total disintegration of those pollutants. An amalgam of
 8 Role of Fungi in Bioremediation and Environmental Sustainability

electrons and electron acceptors can be used to accelerate their metabolism by 60

AU8 microbes. Microbes use organic pollutants for their accumulation during bioreme- 61
diation. Moreover, proliferation of other important nutrients (including nitrogen and 62
phosphorus), as well as minor nutrients (including sulfur and trace elements) occurs 63
(US National Research Council 2000; Qin et al. 2013; Lacerda et al. 2019; Magnin 64
et al. 2019). Microbes also acquire energy by catalyzing energy-efficient chemical 65
reactions that dissociate chemicals from contaminants and transmit electrons 66
(Friesen 2013). These types of reaction are known as oxidation and reduction reac- 67
tions. In natural conditions, it has often been observed that transformation of mol- 68
AU9 ecules and other xenobiotics accompanies their degradation (Fig. 8.1). This process 69
includes phenomena such as co-oxidation, gratuitous metabolism, co-metabolism, 70
and free or accidental metabolism (Tegli et al. 2014; Zengguang et al. 2015). 71
Co-metabolism is a type of metabolism in the presence of an organically active 72
substrate as the primary carbon and energy source, without any nutritional gains. 73
This type of metabolism is a regular microbial activity (Pickering 2000). The meta- 74
AU10 bolic enzymes secreted by bacteria break down the complex organic materials 75
around them to make digestion easier (Segura and Ramos 2013; Kameshwar and 76
AU11 Qin 2019). Such enzymes are usually nonspecific and can function on various types 77
of substrate, including substrate materials that are not beneficial to the bacteria 78
themselves (Ganley et al. 2004; Neifar et al. 2015). 79
Microbes utilize contaminants as sources of carbon for their growth and repro- 80
duction. In this way, they break down the contaminants and transform them into 81
simpler compounds. From this breakdown of contaminants, they obtain energy to 82
reproduce and give rise to new microbial cells. The microorganisms degrade chemi- 83
cal bonds and release electrons, which are then used in production of new microbial 84
cells. When a chemical compound loses electrons, it becomes oxidized, and when it 85
gains electrons, it becomes reduced. This phenomenon is known as a redox reaction, 86

Oxidation,
Recalcitrant compound Reduction,
Intracellular Hydrolysis,
attack:Cytp450 Dehalogenation

Activated Recalcitrant
Extracellular
compound
oxidation
Glutathionylation,
Peroxidase, Conjugation acetylation, methylation
Laccase, Oxidase
Intermediate
Further catabolism
metabolism

Excreted Derivatives Excretion

and storage Vacuole

Fig. 8.2 Mechanism adopted by fungi for bioremediation of toxic, recalcitrant compounds

Fig. 8.1 Bioremediation mechanisms

 A. Tomer et al.

87 where reduction and oxidation occur simultaneously. Most living organisms use
88 oxygen (O2) as an electron acceptor. Thus, we can conclude that organisms degrade
89 organic compounds into simpler molecules such as H2O and CO2 in the presence of
90 O2; this process is known as aerobic respiration. As a result of evolution, some
91 microorganisms do not require O2 to break down chemical compounds (Villela et al.
92 2019). In their processes, contaminants are degraded by nitrate (NO3−) and sulfate
93 (SO42−), and the end products are nitrogen gas (N2), hydrogen sulfide (H2S), and
94 methane (CH4); this process is known as anerobic respiration. The energy released
95 in this process is utilized in cell synthesis. Fermentation is a process in which reac-
96 tions occur in the absence of oxygen, where microbes convert contaminants into
97 simpler by-products such as ethanol, hydrogen, and carbon. In this reaction, the
98 contaminants behave as electron acceptors and electron donors (Cecchi et al. 2019).
99 Some microorganisms convert contaminants into simpler forms that have no AU12
100 beneficial requirements, and this phenomenon is known as secondary utilization.
101 One other phenomenon that occurs simultaneously is co-metabolism, in which the
102 by-products help to detoxify the effects of the reaction. When bacteria are used in
103 degradation of CH4, certain enzymes are produced that degrade the chlorinated sol-
104 vent, which plays no vital role in the growth of the bacteria. Here, the chlorinated
105 solvent serves as a secondary substrate, as it has no role in the maintenance of bacte-
106 rial growth. Another variation due to evolution is reductive dehalogenation, in which
107 halogen atoms in the compound are replaced by hydrogen atoms. Here, detoxifica-
108 tion of the halogen atom occurs, with addition of two electrons to the organic chem-
109 ical compound in the presence of lactate, glucose, and acetate, acting as electron
110 donors. In this reaction, there is no release of energy, but the reaction has a detoxify-
111 ing effect, with removal of a toxic compound, and this is beneficial for production
112 and proliferation of new cells.
113 Nowadays, various bioremediation methods are used to convert toxic organic
114 materials in pesticides, industrial waste, oil spills, etc. into harmless compounds by
115 degrading them. Their transformation into CO2, H2O, N2, hydrochloric acid (HCl),
116 etc. is known as mineralization, and this is the ultimate goal of bioremediation.
117 Heavy metals and radioactive cations are physically removed by phytoremediation
118 or mycoremediation through harvesting of the entire plant or fungus, although they
119 cannot be decomposed (Singh et al. 2014; Ceci et al. 2019). Degrading microorgan-
120 isms obtain carbon, nitrogen, or energy from pesticide molecules (Gan and Koskinen
121 1998). Thus, the most important pesticide degradation process in soil is microbial
122 metabolism (Kearney and Wauchope 1998). Fungi are known to play a vital role in
123 leaf litter degradation; moreover, they are the only organisms on earth that decom-
124 pose wood. Lignin and cellulose are essential components of plant fiber, which is
125 decomposed into humus by extracellular enzymes and acids exuded from fungal
126 mycelia. It is possible to increase the rate of degradation by supplying nitrogen,
127 phosphorus, potassium, and other inorganic elements (Rhodes 2013). Decomposition
128 of starches, celluloses, hemicelluloses, other sugar polymers, and pectins is carried
129 out by molds such as Aspergillus and Botrytis. They are also capable of degrading
130 fats, oils, chitin, and keratin. These molds can be used for biodegradation, in which
131 they degrade paper and textile raw materials such as cotton, linen, and jute. Fungi
8 Role of Fungi in Bioremediation and Environmental Sustainability

such as Mucor thermohyalospora, Cladosporium oxysporum, Phanerochaete chrys- 132

osporium, Trichoderma harzianum, and Aspergillus spp. (e.g., Aspergillus niger 133
and Aspergillus terreus) have the ability to degrade endosulfan, which causes prob- 134
lems in the environment and in living organisms (Bhalerao and Puranik 2007). 135
Fungi can transform pesticides into innocuous substances via certain processes such 136
as esterification, hydroxylation, deoxygenation, and dehydrogenation (Pinto et al. 137
2012; Deng et al. 2015). A few examples are mentioned below. 3-Phenoxybenzoic 138
acid is hydroxylated into 3-hydroxy-5-phenoxybenzoic acid, which is further deox- 139
ygenated into gallic acid and phenol (Deng et al. 2015). A fungal strain of Rhizopus 140
oryzae (CDBB-H-1877) can be used for biosorption of pentachlorophenol through 141
dechlorination and methylation. Aspergillus and Zygomycetes fungi can decolorize 142
and detoxify textile wastewater. Polychlorinated biphenyls (PCBs) are degraded by 143
nonligninolytic enzymes produced by fungi such as Penicillium digitatum, 144
Penicillium chrysogenum, Fusarium solani, and Scedosporium apiospermum 145
(Tigini et al. 2009). White rot fungi are preferred as robust and protective tools in 146
soil bioremediation, as they can tolerate high concentrations of pollutant chemicals 147
(Evans and Hedger 2001; Sasek 2003). Reports have shown that brown rot fungi 148
degrade cellulose, leaving lignin undissolved as brown deposits, while white rot 149
fungi digest lignin, leaving cellulose intact and giving a bleached appearance to 150
wood. Some white rot fungi are also capable of degrading persistent xenobiotic 151
compounds. They include Pleurotus ostreatus, Pleurotus tuber regium, Pleurotus 152
pulmonarius, Agaricus bisporus, Lentinula edodes, Bjerkandera adusta, Irpex lac- 153
teus, and Trametes versicolor (Singh 2006). In addition, these white rot fungi can 154
degrade pesticides, phenols, chlorophenols, polychlorinated biphenyls and dioxins, 155
heavy metals, dyestuffs, and effluent from pulp and paper mills (Singh 2006). They 156
are also capable of degrading environmental pollutants such as CO2, dichlorodiphe- 157
nyltrichloroethane (DDT), lindane, and chlordane. This wide range of activity of 158
white rot fungi is due to (1) production of lignin peroxidase (LiP), manganese per- 159
oxidase (MnP), lactase, and various hydrogen peroxide (H2O2)–producing enzymes 160
(Kirk and Farell 1987; Adenipekun and Lawal 2012); and (2) their mycelial growth 161
habit, which allows rapid colonization of substrates and hyphal extension, enabling 162
penetration of soil to reach pollutants (Reddy and Mathew 2001 and Park et al. 163
2020). LiP and MnP are also produced by Phanerochaete chrysosporium. Studies 164
have shown that Lentinus subnudus has the ability to degrade both metolachlor and 165
heptachlor by up to 94% and atrazine by up to 78% (Nwachukwu and Osuji 2007). 166
Phanerochaete ostreatus has the potential to degrade heptachlor and heptachlor 167
epoxide by up to 89% and 32%, respectively (Purnomo et al. 2014). Aldrin and 168
Dieldrin pesticides can be degraded by Phlebia acanthocystis, Phlebia brevispora, 169
and Phlebia aurea (Xiao et al. 2011). Degradation of effluent from textile industries 170
has been carried out using several hyphomycetes, ascomycetes, and basidiomycetes 171
fungi isolated from marine environments. 172
 A. Tomer et al.

173 8.3 Bioremediation of Contaminated Land

174 Use of microbes for disintegration of contaminants in soil, as well as in water, can
175 be defined as bioremediation. For efficient mycoremediation, it is important to per-
176 form screening to select suitable fungal species that can degrade the relevant con-
177 taminants. Bioremediation can be done using in situ or ex situ approaches (Akcil
178 et al. 2015). The main difference between these two methods is that bioremediation
179 performed on-site as classified as in situ, while bioremediation performed after
180 physical removal of the contaminant substance from the site is classified as ex situ
181 (Margesin et al. 2003). Ex situ treatment for cost-efficient remediation of soil pol-
182 lutants requires chemicals and incineration (Rodriguez et al. 2008; Gillespie and
183 Philp 2013; Mishra and Malik 2014). The main objective of bioremediation is to
184 mineralize pollutants through their transformation intoCO2, H2O, N2, HCl, etc. It is
185 difficult to decompose heavy metals and radioactive ions, as they are converted into
186 less soluble forms. One example is oxidation of uranium(IV) into uranium dioxide
187 (UO2), a less dangerous form that can be removed physically with the help of phy-
188 toremediation or mycoremediation, which may include use of co-cultivation of
189 fungi and plants (Richardson et al. 1992; Graham and Eissenstat 1998; McGrath AU13
AU14
190 and Zhao 2003; Megharaj et al. 2011; Haq and Raj 2020).
AU15
AU16

191 8.4 Bioremediation Potential of Fungi

192 Fungi have been shown to play significant roles in bioremediation of contaminants
193 such as persistent organic pollutants (POPs), textile dyes, coal, paper leather tan- AU17
194 ning effects, pharmaceuticals and personal care products (PPCPs), polycyclic aro-
195 matic hydrocarbons (PAHs), and pesticides (Prasad 2017, 2018). Various reports
196 have described use of fungi from different groups—including Aspergillus,
197 Penicillium, and alkalophilic white-red fungi—for bioremediation and decoloriza-
198 tion of textile dyes, sugar industry effluent, chemicals used in kraft pulp mills, and
199 leather tanning effluent, indicating the diverse substrate choices of these fungi AU18
AU19
200 (Redman et al. 2001; Redman 2002; Rockne and Reddy 2003). Substantial removal
AU20
201 of petrol and diesel contaminants from soil by short-term incubation of Aspergillus
202 niger and Phanerochaete chrysosporium with petroleum hydrocarbons was shown
203 in conjunction with total organic carbon (TOC) elimination, which helps in biore- AU21
204 mediation (Fig. 8.2) (Timmis 2010; Redman and Kim 2011; Echeveria et al. 2020). AU22
 8 Role of Fungi in Bioremediation and Environmental Sustainability

Bioremediation mechanism

Non-Enzymatic Enzymatic
processes process

Biosorption
Biodegradation

Biomineralization
Intracellular Extracellular
Enzymatic Enzymatic
Biodegradation Phase I and phase II Laccases
(cytochrome P450 family Manganese peroxidases
epoxidase and transferases Versatile peroxidases
Stripping
Fig. 8.1 Bioremediation mechanism

AU23 Fig. 8.2 Mechanisms adopted by fungi for bioremediation of toxic, recalcitrant compounds

8.5 Use of Fungal Enzymes in Bioremediation 205

Cellulases, xylanases, amylases, proteases, lipases, laccases, peroxidases, catalases, 206

chitinases, etc. are fungal enzymes with industrial value and can be used in organic 207
AU24 waste management—for example, in organic fractionation (Betancor et al. 2013; 208
Narayanan et al. 2013; Claus 2014). White rot fungi give rise to one or more types 209
of enzyme, depending on the species and environmental conditions. Their role is not 210
limited to degradation of natural lignocellulose substrates; they can also be used in 211
bioremediation requiring degradation of numerous xenobiotic compounds, includ- 212
ing dyes (Nigam 2013). The ligninolytic enzymes secreted by white rot fungi are 213
classified into two categories—MnPs and LiPs—which can be used for lignin oxi- 214
dation in fungal cells. Laccases and certain fungal class II peroxidases produced by 215
white rot basidiomycetes are well known to degrade organic pollutants (Naranjo-­ 216
Briceño et al. 2013; Quintella et al. 2019). 217

8.6 Mycoremediation Using Fungi 218

The name white rot fungi refers to the secretion of enzymes that break down cellu- 219
AU25 lose and lignin, giving the cellulose a white color. About 30% of the bioremediation 220
linked to literature, by means of fungi (Cruz-Hernández et al. 2013; Singh 2005). 221
Bacteria must be adapted for synthesis of specific enzymes that can achieve degra- 222
dation of the relevant pollutant(s). Various organic molecules, including untraceable 223
and persistent components such as PAHs, are susceptible, to differing degrees, to 224
 A. Tomer et al.

225 various strains of the white rot fungi that can degrade them (Egamberdieva and
226 Lugtenberg 2014; Sayyed et al. 2020). Soil polluted with crude oil can be mixed AU26
227 with a lignocellulose substrate—for example, sawdust or maize cob—allowing the
228 fungal species to proliferate in the soil and decompose the crude oil. Moreover,
229 white rot fungi have been shown to effectively disintegrate harmful elements such
230 as dioxins, pesticides, phenols, polychlorinated biphenyls and chlorophenols, efflu- AU27
231 ent, dyestuffs, and heavy metals.

232 8.7 Advanced Technologies Used in Fungal Bioremediation

233 In the field of fungal bioremediation, many technical advances have been made in
234 order to overcome the associated shortcomings. These developments include use of
235 enzymes to reduce the bioremediation time and simplify the process, with greater
236 control over fungal biomass. Bioremediation using immobilized fungi in various
237 bioreactors such as rotating biological contactors and fluidized bed reactors has AU28
238 recently been introduced (Tordoff et al. 2000; Lien et al. 2015; Roccuzzo et al.
239 2020). Bioremediation of benzo[a]pyrene under nutrient-enhanced conditions
240 (involving ligninolysis) results in PAH oxidant monooxygenesis, which was also AU29
241 removed during a subsequent nonligninolytic process (Joutey et al. 2013; Tian et al.
242 2019). Bioremediation of wastewater sludge from sewage treatment plants, mixed
243 with a filament inoculum in a broad-scale bioreactor, has been shown to be sustain-
244 able and environmentally friendly when performed using a continuous process AU30
245 (Connell and Staudigel 2013; Yadav et al. 2019). In a further innovative approach to
246 removal of PAHs by establishing permeable new reactive biobarriers of Trichoderma
247 longibrachiatum on nylon sponges, 90% removal was achieved over a period of AU31
248 14 days (Tyagi et al. 2011; Li et al. 2013).

249 8.8 Bioremediation Using Fungal Cytochromes

250 Fungi have complex enzyme detoxification mechanisms in their bodies for oxida-
251 tive and hydrolyte detoxification. In addition to these structures, some fungi have
252 intracellular xenome networks consisting of cytochrome P450 monooxygenases
253 and glutathione transferases, which enable them to cope with various different pol-
254 lutants. The fungal cytochrome P450 system is a flexible catalyst for region-specific
255 and stereospecific oxidation of nonactivated hydrocarbons. Eradication of pollut-
256 ants can be achieved by use of molecular instruments to generate cytochrome P450
257 monooxygenases quickly and abundantly, including by use of a wide range of yeast
258 expression systems with a viral vector (Arxula adeninivorans) (Tangahu 2011; AU32
259 Singh et al. 2013).
 8 Role of Fungi in Bioremediation and Environmental Sustainability

8.8.1 Factors Affecting Bioremediation 260

The availability of nutrients affects the process of microbial detoxification of con- 261
taminants. Insufficiency of nutrients can directly inhibit the proliferation and 262
enzyme activity of pollutant-degrading organisms. For cell metabolism and efficient 263
proliferation in contaminated environments, microbes require nutrients such as 264
nitrogen, phosphorus, potassium, and minerals (Sagarkar 2013). 265
Environmental conditions such as the pH, temperature, salinity, oxygen level, 266
and availability of water vary from site to site and can inhibit development of the 267
contaminant-degrading microbes that are needed to drive the bioremediation pro- 268
AU33 cess. The pathogens break and grow in the polluted atmosphere, and microbes can 269
metabolize more contaminants under optimal environmental conditions 270
(Egamberdieva and Lugtenberg 2014). 271
AU34 Pollutants may interact with their surroundings to change their bioavailability 272
differs from species to organisms so that great to which the contaminant is free to 273
move into organism can be defined as specified organism. 274
The ability of the microbial community to remove pollutants from a contami- 275
nated site depends on the numbers of microbes at the site and their catabolic effec- 276
tiveness. The presence of soil pathogens can be regulated by both environmental 277
and nutritional factors. 278

8.9 Phytoremediation Using Arbuscular Mycorrhizal Fungi 279

Bioremediation is a method using microbes to treat contaminated soil. In the gen- 280
eral phytoremediation cycle, the combination of AMFs and plants—also known as 281
root–colonizer symbiosis—is involved in soil remediation. AMFs have been found 282
to reduce metal toxicity to plants through a decrease in the rate of root-to-shoot 283
translocation (Fan and Liu 2011). Phytoextraction requires plants that are capable of 284
AU35 storing significant amounts of heavy metals and may further cultivate and discard or 285
remove heavy metals. Organic pollutants (such as PAHs) are transformed by the 286
AU36 microbial activity that is commonly seen around plant roots (Gianinazzi et al. 2010). 287
PAHs are degraded by exudates from plant roots, microorganisms, and AMFs. In a 288
case study in which different methods of soil quality improvement were assessed, 289
AU37 noninoculated soil and soil inoculated with a single AMF mix (indigenous AMFs) 290
were studied (Kumar et al. 2008, 2017; Sim et al. 2019). The AMFs in the soil 291
AU38 inoculated with the indigenous mix were found to be effective in soil quality 292
improvement. The presence of AMF nodules in the soil increased plant growth, 293
AU39 water infiltration, and soil aeration through soil agitation. Phosphorus inoculation in 294
native AMFs is also increased (Francis and Read 1995; Tang 2019). 295
 A. Tomer et al.

296 8.9.1 Fungi as Symbionts

297 In various parts of the world, systematic use of large quantities of fertilizers contain-
298 ing phosphorus has contributed to accumulation of phosphorus in various soil types.
299 Plants that can absorb phosphorus through roots only end AMF growth that clings AU40
300 to their roots by obstructing the fungal flow of sugar and nutrients. AMF hyphae
301 perform two main functions: (1) they serve as a system that absorbs nutrients, and
302 (2) newly formed roots act as plows, breaking the soil hyphal network and hindering
303 its functions (Rodriguez et al. 2009; Mishra et al. 2020). Monoculture of a single
304 crop dominates production of certain fungi that are capable of growing in symbiosis
305 and leads to declines in various other AMFs. Continuous monoculture of a single
306 crop with the same AMF species results in decreased yields. There are a few crops
307 that inhibit root colonization by AMFs (Ruiz-Lozano 2003; Franken 2012), such as
308 Brassica crops, which include spinach, lupin, broccoli, and cauliflower. If Brassica AU41
309 crops are grown in the same rotation, AMF growth in the soil is suppressed.
310 Therefore, an interspersed mixture of supplemental plants is needed to facilitate
311 AMF growth to support AMF numbers in cultivation of these crops. After 8 years of
312 transition from conventional to organic farming in the Farming Systems Trial at the
313 Rodale Institute in the USA, it became clear that larger quantities of fungal spores
314 were produced in organic farming than in traditional farming (Ruiz Sanchez et al.
315 2010; Kumar et al. 2018).

316 8.10 Conclusion

317 Bioremediation is a versatile and environmentally friendly treatment solution and a

318 rapidly growing practice. The capacity of microbes to deal with environmental pol-
319 lutants can be used to disintegrate and/or detoxify them into less harmful forms (US AU42
320 Environmental Protection Agency 1999). Recent research to improve our under-
321 standing of bioremediation mechanisms and genomic developments has shown that
322 whole-genome studies can help to explain and explore bioremediation pathways.
323 Land that is polluted or otherwise unfit for agriculture can be remediated via use of
324 arbuscular mycorrhizal fungi (AMFs) to make it suitable for agriculture. The yield
325 and nutritional value of crops are also increased by use of AMFs. Strong degraders
326 of polycyclic aromatic hydrocarbons are found in AMFs. Because of their sensitiv-
327 ity to a great variety of pollutants, AMFs can also be used in bioassays to test soil
328 and its toxicity levels.
 8 Role of Fungi in Bioremediation and Environmental Sustainability

AU43 References 329

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