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FUNGAL BIOREMEDIATION
Fundamentals and Applications
Editors
Araceli Tomasini
Departamento de Biotecnología
Universidad Autónoma Metropolitana-Iztapalapa
Ciudad de Mexico
Mexico
H. Hugo León-Santiesteban
Departamento de Energía
Universidad Autónoma Metropolitana-Azcapotzalco
Ciudad de Mexico
Mexico
p,
p,
A SCIENCE PUBLISHERS BOOK
A SCIENCE PUBLISHERS BOOK
Cover illustrations reproduced by kind courtesy of Arturo Ruiz Lara and Diana Cuervo Maya
CRC Press
Taylor & Francis Group
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Boca Raton, FL 33487-2742
© 2019 by Taylor & Francis Group, LLC

CRC Press is an imprint of Taylor & Francis Group, an Informa business
No claim to original U.S. Government works
Printed on acid-free paper

Version Date: 20190108
International Standard Book Number-13: 978-1-138-63640-8

9781138636408 (Hardback)
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Library of Congress Cataloging-in-Publication Data

Names: Tomasini, Araceli, editor. | Leon-Santiesteban, H. Hugo (Hector
Hugo), editor.
Title: Fungal bioremediation : fundamentals and applications / editors,
Araceli Tomasini (Departamento de Biotecnologia, Universidad Autonoma
Metropolitana-Iztapalapa, Ciudad de Mexico, Mexico), H. Hugo
Leâon-Santiesteban (Departamento de Energia, Universidad Autonoma
Metropolitana-Azcapotzalco, Ciudad de Mexico, Mexico).
Description: Boca Raton, FL : CRC Press, 2019. | “A science publishers book.”
| Includes bibliographical references and index.
Identifiers: LCCN 2018057729 | ISBN 9781138636408 (hardback)
Subjects: LCSH: Fungal remediation. | Bioremediation. | Factory and trade
waste--Biodegradation.
Classification: LCC TD192.72 .F8425 2019 | DDC 628.5--dc23
LC record available at https://lccn.loc.gov/2018057729
Visit the Taylor & Francis Web site at

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Preface
The large number of people that inhabit our planet and all their activities have caused
severe environmental pollution problems and disruption of the ecological balance.
The air, ground, and water are polluted at different levels depending on the geographic
zone. Some of these pollutants are compounds of natural origin or xenobiotics.
Xenobiotics are human-made compounds for a specific use. Many researchers
have dedicated themselves to studying and understanding the biodegradation of the
pollutants, with the goal of proposing processes for their mitigation. These processes
may be biological and physicochemical; and each of them has advantages and
disadvantages. One of the main problems that prevent institutions and businesses
from applying these processes is that the cost is very high without giving anything
in return economically. Biological processes, also known as bioremediation, are less
expensive to apply than the physicochemical, since they involve the use of living
organisms, such as plants, bacteria, fungi, and/or algae.
We know that nature has her own ways of eliminating organic waste. One of the
main actors involved in these processes are the filamentous fungi. However, the large
number and diversity of compounds that are disposed of into the environment has
overcome the natural capacity of these organisms.
In this book, studies in fungal bioremediation conducted by experts are
mentioned. Studies of fungi bioremediation began since the second half of the last
century. And there have been many advancements in different topics, such as the
isolation of xenobiotic-degrading fungi, degradation mechanisms, enzyme involved
in degradation, molecular studies, proteomic studies, microscopy studies, and
application technologies.
This book shows the newest advancements in the field of fungal bioremediation,
and is mainly addressed to grad students, as well as to professionals and researchers.
The book has three parts: fundamentals, applications, and technologies and tools.
Chapters were written by leading researchers who are doing work on fungi and
bioremediation. In the first part, the characteristics of filamentous fungi and their
role in the environmental remediation are briefly described. The main enzymes that
participate in the processes of degradation of lignin and toxic compounds are also
described. Cutting edge knowledge on fungal enzymes involved in the degradation
of toxic compounds, their production, their activity, and their overexpression are
written in this section.
Each chapter in the second part of the book deals with the degradation and/or
removal of different toxic compounds. Examples of degradation of toxic xenobiotics
are mentioned, which currently cause serious pollution problems, including emerging
iv Fungal Bioremediation: Fundamentals and Applications
pollutants. The degradation of lignin or hydrocarbons are not mentioned, because

both topics are mentioned in all referenced books of fungal bioremediation.
The reader can get information about state-of-the-art advancements and
knowledge in the field of remediation by fungi of chlorophenols, dyes, heavy metals,
emerging pollutants, radioactive compounds, as well as pollutants present in the air.
The chapters mention the most used fungi, the detoxification mechanisms of the
compounds, and the most relevant research conducted till date.
In the third part of the book, tools and cutting-edge technologies used in the
study of the removal and degradation of pollutants by fungi are presented. The first
chapter of this part mentions how proteomics has helped to understand some of the
degradation mechanisms done by fungi. The second one describes genomics as a
very important tool in the understanding of the fungal metabolism, and technological
advancements, such as the use of different bioreactors for the culture of fungi. The
last chapter describes a tool which is not mentioned in other books about these topics,
but which is just as important: microscopy. Topics ranging from the most basic
techniques to the forefront of microscopy are discussed.
We want to thank all contributing authors that put in many hours to the creation
of this book in an enthusiastic and professional manner. Thank you to all the authors
that helped us achieve this project in the best way possible. Finally, we want to thank
our families for their patience, support, and help, which were fundamental for the
creation of this book.
Araceli Tomasini
H. Hugo León-Santiesteban
August 2018
Contents
Preface iii
PART I: FUNDAMENTALS
1. The Role of the Filamentous Fungi in Bioremediation 3
Araceli Tomasini and H. Hugo León-Santiesteban
2. Fungal Peroxidases Mediated Bioremediation of Industrial Pollutants 22
Misha Ali, Qayyum Husain and Hassan Mubarak Ishqi
3. Phenoloxidases of Fungi and Bioremediation 62
Montiel-González Alba Mónica and Marcial Quino Jaime
PART II: APPLICATIONS

4. Chlorophenols Removal by Fungi 93
H. Hugo León-Santiesteban and Araceli Tomasini
5. Azo Dye Decoloration by Fungi 125
M. R. Vernekar, J. S. Gokhale and S. S. Lele
6. Fungal Processes of Interaction with Chromium 173
P. Romo-Rodríguez and J. F. Gutiérrez-Corona
7. Removal of Emerging Pollutants by Fungi: Elimination of Antibiotics 186
Carlos E. Rodríguez-Rodríguez, Juan Carlos Cambronero-Heinrichs,
Wilson Beita-Sandí and J. Esteban Durán
8. Fungi and Remediation of Radionuclide Pollution 238
John Dighton
9. Removal of Gaseous Pollutants from Air by Fungi 264
Alberto Vergara-Fernández, Felipe Scott, Patricio Moreno-Casas and
Sergio Revah
vi Fungal Bioremediation: Fundamentals and Applications
PART III: USEFUL TOOLS

10. A Learning Journey On Toxico-Proteomics: 287
The Neglected Role of Filamentous Fungi in
the Environmental Mitigation of Pentachlorophenol
Celso Martins, Isabel Martins, Tiago Martins, Adélia Varela and
Cristina Silva Pereira
11. Bioreactors and other Technologies used in Fungal Bioprocesses 319
Avinash V. Karpe, Rohan M. Shah, Vijay Dhamale, Snehal Jadhav,
David J. Beale and Enzo A. Palombo
12. Electron Microscopy Techniques Applied to Bioremediation and 354
Biodeterioration Studies with Moulds: State of the Art and
Future Perspectives
Antoni Solé, Maria Àngels Calvo, María José Lora and
Alejandro Sánchez-Chardi
Index 387
PART I
FUNDAMENTALS
CHAPTER-1
The Role of the Filamentous

Fungi in Bioremediation
Araceli Tomasini1,* and H. Hugo León-Santiesteban2
1. Introduction
Environmental pollutants are a serious problem worldwide. They damage the
ecosystems and all forms of life. Examples abound, such as climate change, diseases,
and in some cases, mutagenic alterations that can lead to the death of organisms, just
to name a few.
Pollution problems are increasing due to the rising human population and their
anthropogenic activities. Compounds that cause pollution of the environment are
increasing in quantity and diversity because of the lifestyle of this century that
constantly demands new products and evolving technology. There are many toxic
compounds used as pesticides, biocides, or as energy sources. Others are not toxic, but
their accumulation in the environment is so high, that they can also cause pollution.
Given this problem, many studies have aimed at diminishing the toxic compounds
present in the environment, either in the soil, water, or air. Physical, chemical, and
biological processes have been proposed to remove the toxic compounds. All of them
present advantages and disadvantages and their application depends on the type and
concentration of pollutants and the site where they are present: soil, water, or air.
Biological processes involve organisms or biological activity to degrade or
remove the pollutants; this is known as bioremediation. This chapter will describe
the fundaments of bioremediation by fungi, deal with how they can remove toxic
compounds, and provide some examples.
1
Departamento de Biotecnología, Universidad Autónoma Metropolitana-Iztapalapa, C.P. 09340, Ciudad
de Mexico, Mexico.
2
Departamento de Energía, Universidad Autónoma Metropolitana-Azcapotzalco, C.P. 02200, Ciudad de
Mexico, Mexico.
* Corresponding author: [email protected]
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4 Fungal Bioremediation: Fundamentals and Applications
2. Bioremediation
Many definitions are given for bioremediation. One of the most accepted is:
“bioremediation is a process to clean a polluted site using organisms as plants,
algae, bacteria, and fungi” (Vidali 2001). Another definition used by many authors
is the following: “bioremediation is a biological process to degrade or remove
environmental contaminants from a polluted site, water, soil, and air, using organisms
or biological activity as plants, algae, bacteria, fungi, and enzymes”.
The principal advantage of bioremediation compared to physical and chemical
processes is the cost. In general, it is lower than the physical and chemical
processes, besides that it is a process that does not produce more contaminants. The
disadvantages are that the bioremediation process takes more time than physical or
chemical processes. The second one is that the site to be cleaned must have the right
conditions for the growth and development of organisms that carry out the processes
of removal or degradation of the contaminants.
Bioremediation includes two mechanisms to diminish the polluted compounds,
through degradation involving the chemical modification of the molecules, and
removal, which is a sorption process of the polluting molecules. Removal of toxic
compounds is accomplished through a sorption process, the compounds are absorbed
by a substrate; in this case, a biological substrate, biomass, or agriculture waste that
can be used wet or dry. Biodegradation involves a chemical change of the molecule
to produce other compounds, generally less toxic, and the live organisms or their
enzymatic system are responsible for the compound’s degradation.
Bioremediation involves different organisms. When plants are responsible
for the remediation it is called phytoremediation. This process is one of the most
economic ones, but it takes more time to be accomplished.
Bacteria, fungi, and algae can also be responsible for bioremediation; the
efficiency depends on the type of compounds, the initial concentration of the toxic
compounds, the site to be cleaned, and the microorganism to be used. In many cases,
high efficiency is obtained using a microbial consortium containing a wide diversity
of microorganisms, including different types of bacteria, fungi, and microalgae.
This chapter is focused on studying the role of filamentous fungi in bioremediation
processes; the role of other organisms, such as plants, bacteria, and algae are not
discussed, but it is important to know that bioremediation could also be using some
of these organisms.
3. Filamentous Fungi
Fungi are eukaryotic and heterotrophic organisms that include unicellular and
pluricellular fungi. The unicellular are yeasts, and the pluricellular are represented
by all types of filamentous fungi, called as such due to their way of growth. Yeast are
anaerobes and aerobes, and filamentous fungi are exclusively aerobes. This chapter
will deal only with filamentous fungi.
They develop a tubular structure named hypha, the hyphae form the mycelium,
fungi can reproduce by sexual or asexual spores, and by vegetative means from
a mycelium segment. There are many types of filamentous fungi; they exist in
terrestrial and aquatic habitats.
The Role of the Filamentous Fungi in Bioremediation 5
These fungi can grow as saprotrophs, which obtain their nutrients from dead
organisms. They can grow also in symbiosis, which means a common life between
two organisms including parasitism and mutualism. Parasitism is when only one
of the organisms benefits and mutualism when both organisms are benefited; for
example, mycorrhizae (mutualism with roots of plants) or lichens (mutualisms with
algae) (Carlile et al. 2001).
They can grow in soil producing many hyphae that spread profusely all over the
ground; despite fungi being microscopic, hyphae of Armillaria ostoyae can reach a
spread rate of 1 m year–1 (Peet et al. 1996). Armillaria bulbosa, a filamentous fungus
type, has been reported as one of the largest and oldest-living organisms (Smith
et al. 1992).
Some fungi are pathogenic agents of plants and they are responsible for the loss
of agricultural products, which are an important food source. However, many other
filamentous fungi are used to produce secondary metabolites, such as antibiotics,
immunosuppressors, anticholesterolemics, etc. (Barrios-Gonzalez et al. 2003).
An important role to note is that fungi are the primary organisms responsible for
degrading organic matter in nature.
Filamentous fungi grow in soil, and many of them degrade organic compounds
present in dead organisms, both animal and vegetal. Organic matter is converted into
small molecules that can be used by other organisms as a source of energy, carbon,
and nitrogen. The ability of fungi to degrade organic compounds has been exceeded
by the large number of organic compounds present in the soil due to the increase of
human populations in our planet. This ability has also been affected by the presence
of toxic compounds, such as xenobiotics. These last compounds are not present
in nature, but they are produced by humans for specific uses, such as pesticides,
biocides, or are the result of waste or byproducts from industrial processes.
Bioremediation by fungi is an alternative process to remove, degrade, or render
harmless toxic compounds using natural biological activity. The fungi belonging to
the basidiomycetes were the first assayed to degrade toxic compounds, as it was
observed that this class of fungi grows on fragments of trees that are lying on the
ground in forests, causing their rotting. Depending on the type of rotting that they
cause on wood, fungi are called white-, brown-, or bland-rot fungi. These fungi have
a common characteristic, the production of an enzymatic system able to degrade
lignin. For this reason, they are also named ligninolytic fungi. Phanerochatete
chrysosporium was one of the first white-rot fungi studied, showing its ability to
degrade lignin, toxic compounds, and its relationship with the enzymatic system
(Bumpus et al. 1985).
From the 1980s onwards, many fungi have been studied and found, which can be
used in environmental bioremediation. It has been shown that ligninolytic fungi could
degrade diverse types of toxic compounds, such as hydrocarbons, chlorophenols,
polychlorinated biphenyls, etc., because of their enzymatic system.
Basidiomycetes, such as Phanerochaete, Trametes, Pleurotus, Lentinus, among
others, can degrade toxic compounds (Table 1.1). Many of them have been isolated
from natural sources, mainly from the soil or wastewater contaminated with toxic
compounds (Singh 2006, Fackler et al. 2007, Harms et al. 2011, Seigle-Murandi
et al. 1993, Lee et al. 2014, Mineki et al. 2015).
6
Table 1.1 Filamentous fungi used in bioremediation.
Class Fungi Use References

Basidiomycete Phanerochaete chrysosporium Pentachlorophenol and anthracene degradation Mileski et al. (1988), Mohammadi et al. (2010)
P. chrysosporium Dichlorophenol degradation Huang et al. (2017)
P. chrysosporium Blue 4 dye sorption Bayramoğlu et al. (2006)
Phanerochaete velutina Wood decomposing Darrah and Fricker (2014)
Phanerochaete sordida Dioxin degradation Sato et al. (2003)
Bjerkandera adusta Pentachlorophenol degradation Rubilar et al. (2007)
Trametes versicolor Dye decolorization BTEX* oxidation Libra et al. (2003), Aranda et al. (2010)
Trametes modesta Trinitrotoluene degradation Nyanhongo et al. (2006)
T. versicolor Sulfonamide degradation Rodríguez-Rodríguez et al. (2012)
Pleurotus ostreatus Olive cake degradation Saavedra et al. (2006)
Pleurotus pulmonarius Aromatic hydrocarbons D’Annibale et al. (2005)
P. ostreatus Ni, Cr(VI), Zn, Cd sorption Javaid et al. (2011)
P. ostreatus Naphthalen sulphonic acid polymers degradation Palli et al. (2016)
B. adusta Naphthalene sulphonic acid polymers degradation Palli et al. (2016)
Lentinula edodoes Phenol degradation Ranjini and Padmavathi (2012)
Irpex lacteus Soil bioremediation Novotný et al. (2000)
Panus tigrinus Chlorophenol degradation Leontievsky et al. (2002)
Anthracophyllum discolor Pentachlorophenol degradation Rubilar et al. (2007), Cea et al. (2010)
Fungal Bioremediation: Fundamentals and Applications
Geotrichum sp. Azo dyes biotransformation Máximo et al. (2003)

Gloeophyllum striatum Fluorophenol degradation Kramer et al. (2004)
Daldinia concentrica Dibutylphthalate degradation Lee et al. (2004)
Agaricus augustus Tribromophenol degradation Donoso et al. (2008)
Agaricus bisporus PAH and Pb removal García-Delgado et al. (2015)
Funalia trogii Dyes decolorization Özsoy et al. (2005)
Phlebia tremellosa Lignin degradation Fackler et al. (2007)
Oxyporus latemarginatus Lignin degradation Fackler et al. (2007)
Lentinus tigrinus Phenol degradation Kadimaliev et al. (2011)
Eisenia fetida Olive cake transformation Saavedra et al. (2006)
T. versiclor Naproxen degradation Marco-Urrea et al. (2010)
Pycnoporus sanguineus Metal sorption Zulfadhly et al. (2001)
P. sanguineus Cu(II) sorption Yahaya et al. (2009)
Hyphomycete aquatic Clavariopsis aquatica Nonylphenol degradation Junghanns et al. (2005)
Ascomycete Aspergillus spp. Banana waste degradation Shah et al. (2005)
Asperigllus niger var tubingensis Cr(VI) Sorption/reduction Coreño-Alonso et al. (2009), (2014)
Asperigllus niger Pentachlorophenol sorption Mathialagan and Viraraghavan (2009)
Aspergillus awamori Phenol, catechol, dichlorophenol, dimethoxyphenol Stoilova et al. (2006)
degradation
Aspergillus fumigatus Anthracene degradation Ye et al. (2011)
A. fumigatus Cd, Cu, Ni, Pb and Zn accumulation Dey et al. (2016)
Aspergillus spp. Cr and Cd bioaccumulation Zafar et al. (2007)
Aspergillus oruzae Anthraquinone bisorption Zhang et al. (2015)
Phylosticta spp. Banana waste degradation Shah et al. (2005)
Trichoderma virgatum Pentachlorophenol degradation Cserjesi and Johnson (1972)
Gloeophylhum striatum Dichlorophenol and pentachlorophenol degradation Fahr et al. (1999)
Trichoderma longibrachiatum Pentachlorophenol degradation Carvalho et al. (2009)
Trichoderma spp. Soil bioremediation Harman et al. (2004),
Ezzi and Lynch (2005)
Trichoderma spp. Cyanide degradation Ezzi and Lynch (2005)
Trichoderma harzianum Soil bioremediation Matsubara et al. (2006)
Table 1.1 contd. ...

8
...Table 1.1 contd.
Class Fungi Use References

Ascomycete Fusarium spp. Cyanide degradation Ezzi and Lynch (2005)
Fusarium solani DDT degradation Mitra et al. (2001)
Fusarium oxysporum Phthalate degradation Kim et al. (2003)
Fusarium spp. Zn sorption Velmurugan et al. (2010)
F. solani Anthracene and benzl[a]anthracene degradation Wu et al. (2010)
Penicillium janczewskii Pentachlorophenol degradation Carvalho et al. (2009)
Penicillium restictum Black 5 dye sorption Iscen et al. (2007)
Cordyceps sinensis Dioxane degradation Nakamiya et al. (2005)
Talaromyces helicus Biphenyl degradation Romero et al. (2005)
Byssochlamys nivea Pentachlorophenol degradation Bosso et al. (2015)
Scopulariopsis brumptii Pentachlorophenol degradation Bosso et al. (2015)
Zygomycete zygomycetes Vanillic acid transformation Seigle-Murandi et al. (1992)
Cunninghamella echinulata Pentachlorophenol degradation Seigle-Murandi et al. (1993)
Rhizopus oryzae Pentachlorophenol degradation León-Santiesteban et al. (2014), (2016)
Rhizopus spp. Cr and Cd bio-accumulation Zafar et al. (2007)
Rhizopus nigricans Cr(VI) sorption Bai and Abraham (2003)
Rhizopus sp. Arsenic sorption Jaiswal et al. (2018)
Amylomyces rouxii Pentachlorophenol degradation Montiel et al. (2004), Marcial et al. (2006)
Mucor rouxii Pb, Ni, Cd and Zn sorption Yan and Viraraghavan (2003)
Fungal Bioremediation: Fundamentals and Applications
Mucor plumbeus Pentachlorophenol degradation Carvalho et al. (2011), (2013)

Mucor ramosissimus Pentachlorophenol degradation Szewczyk et al. (2003),
Szewczyk and Dlugoński (2009)
Mucor hiemalis Ni(II) sorption Shroff and Vaidya (2011)
Mucor hiemalis Cr(VI) Tewari et al. (2005)
Mucor sp. Cd and Pb sorption and accumulation Deng et al. (2011)
*benzene, toluene, ethyl benzene, and xylene
4. Biosorption
Removal of toxic compounds through sorption by fungal biomass is one of the
bioremediation processes. The compounds are not modified, they are only removed
or eliminated from the polluted site and concentrated into a support, and this is called
sorption. If the support is biological, the process is called biosorption.
Sorption is a physicochemical process where the molecules from a fluid phase,
also called sorbate, are adhered to a surface of a solid, the sorbent. Adsorption
processes are useful tools to remove chemicals from polluted sites, adsorption
technologies can use inorganic and organic sorbents and they may be physical or
chemical. In physical sorption, the involved forces are weak, such as van der Waals
interactions, and electrostatic interactions formed by gradient-dipole or -quadruple
interactions. Chemical sorption is due to the formation of a chemical bond between
the sorbate and the sorbent. These interaction forces are stronger and more specific
than the forces in physical sorption.
The main advantage of biosorption is that sorbents are cheap and biodegradable;
they include agricultural wastes, such as wood chips, sugar cane bagasse, wheat
shell, microbial biomass from bacteria, fungi, yeasts, and microalgae, etc. Fungal
biomass has been studied to remove toxic compounds from polluted water. It was
applied first to the removal of heavy metals, because fungal biomass is efficient to
remove these ions. Biosorption occurs when the metal is fixed to the inactive or non-
viable biomass, wet or dry.
Biosorption can occur in different parts of the cell, mainly in the cell wall by
mechanisms, such as ionic exchange, chelation, or formation of a complex. Fungal
biomass waste from the fermentation industries could be used as sorbent (Kuyucak
and Volesky 1988, Volesky 1990). Biosorption of toxic compounds or heavy metals
by fungal biomass is due mainly to the cell wall of fungi, which contains between
80 and 90% of polysaccharides, mainly chitin, glucan, mannan, galactosan, and
chitosan, in some cases, cellulose. The rest, 10–20%, is constituted by proteins,
lipids, polyphosphates, and inorganic ions that constitute the cement of the cell wall.
The specific composition depends on the fungus and the identity and quantity of the
different functional groups of the fungal biomass. Sorption ability of fungi is related
to the functional groups present in the cell wall.
Metals can be removed also by living biomass involving metabolic processes;
in this case the process is named bioaccumulation. Removal of heavy metals by
bioaccumulation and biosorption by fungi has been studied for approximately 40
years. Filamentous fungi, such as Aspergillus sp., Rhizopus sp., Mucor rouxii,
Penicillium sp., Trametes sp., among others, have been widely used to remove heavy
metals, including As, Cr, Ni, Pb, Cd, Zn, Hg, Al, etc.
Mechanisms used by fungi to remove metals from the environment have been
reported (Tsezos et al. 1997, Coreño-Alonso et al. 2014, Awasthi et al. 2017).
Biosorption of metal ions is due to the physicochemical interactions between
the functional groups of the fungal cell wall and the metal ions by mechanisms,
such as physical sorption, ion exchange, and chemical sorption, such as chelation,
complexation, and coordination. This sorption can be reversible, which means the
metal can be desorbed from the biomass. Metal bioaccumulation by fungi involves
cell metabolism; the metal must pass through the cell membrane into the cytoplasm
and this action has been related to different defense mechanisms of fungi against
toxicity caused by metals. In some cases, it can be metal precipitation or extracellular
bioaccumulation, involving both mentioned mechanisms (Sağ 2001, Dhankhar and
Hooda 2011). The characteristics of fungal biomass have allowed adsorbing many
types of toxic compounds besides metals.
The fungal biosorption approach is an alternative technique to remove different
types of pollutants, such as phenolic compounds and dyes (Crini 2006, Kumar and
Min 2011). The type of biomass and their previous treatment, type of pollutant,
initial concentration of the compound, biomass concentration, pH, and temperature
influence the process.
Fungal biomass is washed with water and it can be used either wet or dry. In
some cases, the biomass is pre-treated to modify the functional groups of the fungal
biomass, it can also be used immobilized, in both cases, the modification is made
to increase the sorption ability. pH plays an important role because the solubility
and ionization of toxic compounds depend largely on the pH of the phase where
the toxicants are in the environment, mainly if the pollutants are in aqueous phase
(Aksu 2005, León-Santiestebán et al. 2011, Kumar and Min 2011).
Fungal biosorption has been studied for many years; however, its application
is limited mainly because more basic knowledge is needed on aspects such as
physicochemical conditions, effect of parameters, and because the technology has
not been fully developed yet.
5. Biodegradation
The degradation process is the chemical modification of the toxic compounds into
less toxic or innocuous molecules. The mineralization process refers to the complete
degradation of the toxic compound until CO2 and water are obtained. However,
mineralization is not always achieved, the most common is a partial degradation,
in this case, some metabolites derived from the toxic compounds are produced.
Generally, these metabolites are less toxic or completely innocuous; however, in a
few cases the metabolites produced could be even more toxic than the original.
Fungi can degrade a vast variety of toxic compounds due to their enzymatic
system. White rot-fungi degrade lignin, the most abundant resistant biopolymer in
nature, have shown the ability to degrade toxic and xenobiotic compounds, using the
same mechanisms that they use to degrade lignin. Phanerochaete chrysosporiun is
one of the first and most studied fungus to degrade toxic compounds, its study began
in the 1970s and, since then, many studies on the ability to degrade toxicants with this
fungus have been reported (Mileski et al. 1988, Bogan and Lamar 1995, Laugero et
al. 1997, Yadav et al. 1995). Table 1.1 shows some of the class and type of fungi used
in bioremediation. These fungi have the characteristic to produce oxidase enzymes,
which are involved in the degradation of the recalcitrant and xenobiotic toxicants,
such as polycyclic aromatic compounds (PAHs), chlorophenols, polychlorinated
biphenyls, synthetic polymers, dyes, pesticides, benzene, toluene, ethyl benzene, and
xylene (BETX), chlorinated monoaromatics, halogenated organic compounds, and
others. These compounds can be degraded by different fungi, mainly basidiomycetes,
such as the white-rot fungi species that include Phanerochaete, Bjerkandera,

Trametes, Pleurotus, Phlebia, and Ceriporipsis (Cameron et al. 2000), and brown-
rot fungi, for example Lentinus, Lentinulla, and Gloeophyllum. Fungi different
from basidiomycetes have also been studied regarding their ability to degrade toxic
compounds.
Degradation of polycyclic aromatic hydrocarbons (PAHs) by fungi has been
reported by many authors. Most studies showed that fungi degrade this group of
toxicants by co-metabolism, which means fungi cannot use the PHAs as the sole
source of carbon, as accomplished by bacteria (Ghosal et al. 2016). Ligninolytic
and non-ligninolytic fungi can degrade PHAs, however ligninolytic fungi have been
studied more and present some advantages.
The metabolic pathway of degradation of PHAs by non-ligninolytic fungus
is different from the degradation pathway used by ligninolytic fungi. Degradation
of PHAs by non-ligninolytic fungi, involving cytochrome P450, can produce
dihydrodiols and epoxides that are more toxic than the original PHAs. Ligninolytic
fungi degrade PHAs by peroxidases and phenoloxidases producing quinones,
i.e., less toxic molecules (Hammel 1995, Tortella et al. 2005, Cerniglia and
Sutherland 2010, Treu and Falandysz 2017). Co-metabolic anthracene and pyrene
degradation by A. fumigatus and Pseudotrametes giboosas, respectively, was
reported by Ye et al. (2011) and Wen et al. (2011), the last authors indicated that the
degradation rate is high in co-metabolic conditions.
Romero et al. (2005) reported biotransformation of biphenyls by an ascomycete of
the genus Talaromyces, finding that its oxidation produces dehydroxylate derivatives
and their glucosyl conjugates. Pentachlorophenol degradation by zygomycetes, such
as Mucor, Amylomyces, and Rhizopus has been reported by Carvalho et al. (2011),
Montiel et al. (2004), and León-Santiesteban et al. (2014). Table 1.1 depicts other
examples of toxicants’ degradation by different filamentous fungi.
There are diverse mechanisms of toxic compound degradation by fungi, for
example, dehalogenation, which could be through oxidative or reductive reactions,
methylation, hydroxylation, conjugation, breaking of the aromatic rings, and
polymerization are the most common reactions. The mechanism to degrade a toxicant
by fungi depends on the fungi, the characteristic of toxic compounds, the conditions of
the reaction, the composition of the culture media. For example, if fungi are grown in
nitrogen-deficient cultures, or with glucose or another carbon source, the mechanism
to degrade a specific toxicant would be different. León-Santiesteban et al. (2014)
reported that in PCP degradation by R. oryzae ENHE, nine metabolites were found
when the fungus grew with glucose-ammonium sulfate as source of carbon and
nitrogen. When the fungus grew in glutamic acid-sodium nitrate as source of carbon
and nitrogen, only three metabolites from PCP degradation were found.
Szewczyl et al. (2018) propose a biodegradation pathway of the herbicide
ametryn, a herbicide from the triazines group, by Metarhizium brunneum. This
fungus is an entomopathogen fungus tested to infect insects. These authors showed
the ability to degrade ametryn and reported the metabolites produced from the
degradation process.
As mentioned before, a great diversity of toxicants has been degraded by fungi,
and in some cases, the mechanisms of degradation have been elucidated. We mention
only some examples, there is much information about the degradation by fungi of
toxic compounds including pesticides, PHAs, halogenated compounds, and emergent
pollutants, such as pharmaceutical and personal care products (Table 1.1). Authors
have focused the study on toxic degradation by fungi using different tools, such as
fungi isolation, biochemistry, molecular biology, and analytical chemistry in order to
understand and improve fungal bioremediation. Regardless to the focus of the study,
the goal is to propose efficient processes of toxicants’ degradation by fungi.
Currently, the enzymatic, redox reactions, and degradation pathways are known,
as well as the metabolites produced from some toxic compounds degradation, such
as PHAs, chlorophenols, azo dyes, by fungi. However, the degradation pathways
by fungi of many recalcitrant compounds, such as emergent pollutants, are not yet
known, just as the role of enzymes involved in degradation is not understood in
many cases.
Molecular biology studies of fungi have been helping to understand and increase
the ability of these microorganisms to degrade toxicants (Bogan et al. 1996, Huang
et al. 2018, Szewczyk et al. 2018). Many studies about genetic improvement of fungal
strains for degradation of toxic compounds have been reported (Mitra et al. 2001,
Gao et al. 2010, Tripathi et al. 2013). For instance, Syed et al. (2010) reported the
identification and functional characterization of P450 monooxygenase involved in
the oxidation of PAHs by P. chrysosporium using a genome-to-function strategy.
Molecular technologies have allowed better understanding of the role of Ascomycota
fungi in the transformation of toxicants (Aranda 2016).
In addition, some other applications have been proposed to remove toxic
compounds from polluted sites. Bioaugmentation or biostimulation is referred to
increase the presence of fungi responsible for or involved in degradation processes
of some compounds in situ. The above mentioned can be made first by isolating
fungi from the site, then culturing them in the laboratory and, finally, inoculating the
polluted soil with the cultured fungi. Lladó et al. (2013) reported the use of two fungi,
Trametes versocilor and Lentinus tigrinus, for PAH removal from contaminated soil.
The authors reported that PAH fractions remain in the soil contaminated by creosote
after 180 days and that the biostimulation of the soil increases degradation rates.
Degradation of anthracene by bioaugmentation of T. versicolor in a composting
process was reported by Sayara et al. (2011).
While the use and the ability of fungi to degrade toxic compounds have been
studied extensively, more studies are still needed for the scaling-up of the process.
In situ bioremediation is still the objective to achieve.
6. Enzymes
The chemical modification of the toxicant is carried out by enzymatic reactions,
involving oxidoreductase enzymes. This class of enzymes catalyzes redox reactions
with transfer of hydrogen and oxygen atoms or electrons among molecules.
Oxidoreductases include dehydrogenases, oxidases, peroxidases, and oxygenases.
Table 1.2 shows the mechanisms of actions of these enzymes.
Peroxidases and phenoloxidases are the main enzymes produced by fungi.
Peroxidases are hemoproteins and the hydrogen peroxide is necessary as an electron
acceptor. Phenoloxidases are enzymes containing copper that catalyze the oxidation
of phenolic compounds without cofactor, using molecular oxygen for the reaction.
Table 1.2 Mechanisms of action of oxidoreductase enzymes.
Enzyme Mechanism of action

Oxygenases Transfer oxygen atoms from diatomic oxygen
Oxidases Transfer electrons to diatomic oxygen
Peroxidases Transfer electrons to hydrogen peroxide
Dehydrogenases Transfer hydride ions
Some of these enzymes were found in the culture broth of P. chrysosporium, and
it was shown that their activity is dependent on H2O2 (Forney et al. 1982, Paszczyński
et al. 1985, Hatakka 1994), and they were named lignin peroxidases by Tien and
Kirk (1988). It was shown that the enzymatic system produced by P. chysosporium
and other basidiomycetes can degrade lignin; these enzymes are called ligninolytic.
Lignin peroxidase and manganese peroxidase use hydrogen peroxide produced by
glyoxal oxidase or aryl alcohol oxidase produced by the same fungi. Laccases use
the four copper atoms to catalyze the electron reduction of molecular oxygen to
water (Kudanga et al. 2011, Sinsabaugh 2010). The main feature of these enzymes
is that they are non-specific; thus, they can degrade diverse substrates, including
toxic compounds. Many authors have studied fungal ligninolytic enzymes and their
activity and ability to degrade recalcitrant compounds, such as BTX, polyaromatic
hydrocarbons (PHA), polychlorinated biphenyls, etc.
The most studied fungal enzymes to degrade toxic compounds are lignin and
manganese peroxidases, as well as the phenoloxidases, laccase, and tyrosinase.
These enzymes are mainly produced and excreted by basidiomycetes, such as
P. chrysosoporium, Trametes, Pleurotus, Pycnoporus, etc. Other fungi, such as
Ascomycetes that include Aspergillus, Penicillium, Trichoderma, and Zygomycetes
such as Mucor, Rhizopus, as well as Amylomyces, also produce peroxidases and
phenoloxidases (Durán and Esposito 2000, Montiel et al. 2004). Other enzymes, such
as versatile peroxidases, reductive dehalogenases, cytocrome P450, monooxygenases,
and peroxygenases, produced by fungi are involved in the degradation process
of toxicants (Hofrichter et al. 2010, Subramanian and Yadav 2009, Reddy and
Gold 1999, McErlean et al. 2006, Syed et al. 2010). P. chrysosporium, P. ostreatus,
and B. adusta showed ability to degrade PHAs and the enzymes produced by these
fungi involved in the degradation pathways have been reported (Kadri et al. 2017).
Molecular studies aimed at increasing the enzymatic activity produced by fungi
have also been published. There are works reporting the regulation and expression
of peroxidases and phenoloxidases, heterologous expression, transcriptome and
proteomic studies of fungi producing these enzymes (Kellner and Vandenbol 2010,
Piscitelli et al. 2010, Manavalan et al. 2011, Huang et al. 2011, Huang et al. 2017,
Carvalho et al. 2013, Janusz et al. 2013, Montiel-González et al. 2009).
Immobilization of enzymes or of the enzyme producing microorganism used in
biodegradation processes has been proposed with the advantage that it could be used
several times and the degradation process would be environmentally friendly (León-
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Santiestebán et al. 2011, Voběrková et al. 2018). Immobilization of enzyme could be

accomplished on inert supports such as nylon fibers or on organic ones such as sugar
cane bagasse (León-Santiestebán et al. 2011, Mohammadi et al. 2010). Yang et al.
(2017) reported the degradation of antibiotics by immobilized laccase at pH 7.0 in 48
hours, removing approximately 60% of tetracycline and oxytetracycline.
Enzymes play a key role in the degradation processes and have been widely
studied; the next two chapters deal with these fungal enzymes.
Conclusions
The role of fungi as natural degraders of organic material is very important in
the environment, although their capacity has been exceeded by the amount and
the diversity of compounds. The interest of using fungi to degrade toxicants has
increased, due to the current pollution problem and to the low cost that their use
may imply.
To understand how fungi can degrade toxicants and increase their ability,
physiological, biochemical, and molecular studies, including genomics,
transcriptomics, and proteomics, have been made. All this effort has generated a
very broad knowledge of fungi, revealing their potential in bioremediation processes
to eliminate hazardous compounds.
However, for a practical application, bioremediation in situ, more studies are
needed to know the technology that could be used. Bioaugmentation is not an efficient
technology, so far, because of the competition among native microorganisms.
Another drawback is the control of the environmental conditions required for fungi
growth. The addition of enzymes to the site is complicated, since the enzymatic
activity depends on parameters, such as pH, humidity, and temperature, which are
not possible to control in situ. There are some studies reporting the scaling-up from
laboratory to field applications; however, it is necessary to study this subject more to
propose potential soil bioremediation processes using fungi.
Studies and financial investment should be directed to implement technologies
applied in situ, which will help and increase the ability of fungi to degrade hazardous
compounds, for the bioremediation of polluted environments. It should be emphasized
that although bioremediation is a very slow process compared to chemical processes,
it is not expensive and is friendly for the environment.
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of the outside world set down in the midst of military activities. There
mother, wife, sister, sweetheart, friend could meet her soldier or
sailor lad, could spend the night if necessary and have good,
inexpensive meals. It was the scene of many impromptu weddings,
the hostess of the house and her assistants taking charge of the
arrangements, when lovers decided suddenly to be married before
the ocean and the chances of battle should separate them. The Y.
W. C. A. carried its work to France, and in its Hostess Houses there
looked after the welfare of the women workers for the American
Expeditionary Force and its canteens followed the American troops
even to north Russia, where they were established in Murmansk and
Archangel.
All of the great religious bodies of the country joined at once in the
effort to lessen for the army and navy men the hardships of war, to
surround them with as many as possible of the comforts of civilized
life and to uphold them physically, mentally and morally. People of
Protestant faith gave their support mainly to the long established and
widely reaching organization of the Y. M. C. A., members of the
Catholic Church, working through the National Catholic War Council,
supported the endeavors of the Knights of Columbus, and the
Jewish Welfare Board, with American Jewry behind it, turned its
attention especially to soldiers of that faith. And the Salvation Army,
with its years of experience in caring for the needs of humanity and
upholding morale, was early in the field. All these organizations
coöperated in the most cordial way, supplementing one another’s
effort and joining their endeavors whenever the best results could be
gained in that way, two or more of them sometimes using the same
building. The friendly hand, the good cheer, the comforts each had to
offer were ready for any man in uniform without a thought as to his
religious affiliations. Each held its religious ministrations in reserve
for those who asked for them and, for the rest, based its abundant
and many-sided service solely on the desire to help the American
Army fight the battle of justice and liberty. Their one purpose was to
big-brother the fighting forces of the nation and, whether in training
camp or debarkation port, on transport or battleship, behind the lines
in France or at the very front, to be ready with whatever help and
cheer and comfort it was in their power to give when it was wanted.
In a Red Triangle Hut in the Battle Zone
The Jewish Welfare Board was the youngest of all these
organizations, having been formed after our entrance into the conflict
for the purpose of helping to win the war by carrying out the policies
of the War Department with regard to the welfare and the morale of
the soldiers. Behind it were three and a half million citizens of the
Jewish faith and, while it functioned on its religious side for the
benefit of the 175,000 men of the Jewish religion in the Army and
Navy, in all its other activities it was nonsectarian and worked as
generously and cordially for one as for another. In the training camps
of the Army and the Navy in the United States it had many huts and
nearly three hundred field workers who arranged entertainments,
classes and study groups, provided religious services, and taught
the English language and the principles of American citizenship to
men new to America. In two hundred communities near training
stations the representatives of the Welfare Board coöperated with
the War Camp Community Service in all the phases of its activities.
Overseas it had headquarters in Paris and at the end of hostilities it
was preparing to establish others at debarkation ports and in cities
near the large camps of the A. E. F. and was ready to send a
hundred men and women workers to take charge of them. Its club
rooms in Paris were equipped with books, music, games and other
means of social enjoyment and the organization, by coöperation with
a French society, arranged to have Jewish soldiers entertained in
French homes of their own faith. Through the suggestion of the
Welfare Board a number of rabbis were commissioned as chaplains
with the fighting forces, each of them being provided with a monthly
allowance to expend upon small comforts for his boys. They held
Jewish holyday services back of and almost in the front line
trenches, in cities and villages, once in the ruins of a Roman Catholic
Cathedral and again in a large Y. M. C. A. hut. At one service, at
which the rabbi, coming from another sector, arrived a little late, he
found that the local Knights of Columbus Chaplain had kept the
meeting together for him and opened it with a preliminary prayer.
The National Catholic War Council, organized to direct the war-
aiding activities of all Catholic forces, operated a million-dollar chain
of Visitors’ Houses at army and navy training camps and of service
clubs in communities and embarkation ports, where it worked in
coöperation with the War Camp Community Service. Under its
supervision was the society of the Knights of Columbus which, at the
close of hostilities, had in the United States several hundred
buildings and 700 secretaries and overseas more than a hundred
buildings and huts, with many more in preparation, and over 900
workers. It had service clubs in London and Paris which provided
reading, lounging and sleeping rooms, and all such club comforts,
while its huts behind the lines furnished centers of comfort, cheer,
entertainment and small services of many sorts. It operated a great
fleet of motor trucks which carried supplies up to the firing line and
into the front trenches. Nothing was more welcome to the battle-
weary soldiers relieved from front line duty than these “K. C.” rolling
canteens with their hot drinks, cigarettes and other comforts. The
organization shipped to the other side and gave to soldiers and
sailors many tons of supplies, including cigarettes by the hundreds of
millions and huge amounts of chewing gum, soap, towels, stationery,
candy and chocolate. It had more than a hundred voluntary
chaplains on service with the troops, many of whom carried money
furnished by the society to aid in providing comforts for the welfare of
the soldiers.
The Salvation Army won a peculiar place in the hearts of our
fighting men by the simple hominess and complete self-abnegation
of its service. Its huts and hostels were in all the important training
camps at home, while overseas the Salvation Army uniform in some
kind of a structure or dugout welcomed the army lad in the big camp
areas, in the supporting lines and in the forward troop movements up
to the rear of the front line forces and trenches. It had overseas more
than 1200 officers, men and women, operating 500 huts of one sort
or another, rest rooms and hostels. It had forty chaplains serving
under Government appointment and it supplied nearly fifty
ambulances. Its method was to put a husband and wife in charge of
a canteen or hut, the man making himself useful in any way that
offered, the woman making doughnuts and pies, chocolate and
coffee for the ever hungry doughboys, and doing for them whatever
small motherly service was possible. In their huts the men could
always find warmth and light and good cheer, music and games and
good things to eat that were touchingly reminiscent of boyhood and
home. Shells screamed overhead, gas floated back from the front
and the earth shook with the roar of battle, but the Salvation Army
workers stood to their self-imposed duties regardless of their own
comfort or danger and had ready for the long lines of soldiers coming
and going a smiling, heartfelt welcome and huge quantities of pies
and doughnuts and hot drinks. Its canteens were always open, day
and night, and none of its workers was sent overseas without special
training.
By far the largest, oldest and most important of these welfare
organizations was the Young Men’s Christian Association, which
expanded a total of nearly $80,000,000 on a system of war service
so vast that the sun was rising upon it through every hour of the day.
Within a few hours after the United States entered the war the Y. M.
C. A. offered its entire resources to the Government. At the end of
hostilities it had overseas over 7,000 workers, of whom 1,600 were
women; in the American Expeditionary Forces it had 1,900 war
service centers, nearly 1,500 in the French armies, several hundred
in Italy, with more in Russia and Siberia; in the United States it had
950 of these centers and 6,000 workers and it was represented in
every cantonment and training camp for Army or Navy from end to
end of the country. On this side, it paid for its huts and their
equipment a total of more than $6,000,000, while overseas the
similar expenditure went beyond $5,000,000, making a total of well
over $11,000,000 invested in the equipment with which to give our
soldiers and sailors rest and cheer, entertainment and comfort. The
cost of the operation of these centers amounted, for the duration of
the war, to over $6,550,000.
In the home camps and cantonments the “Y” centers had an
average of nearly 20,000,000 visits from soldiers and sailors per
month, while in them at the same time were written letters on free
Red Triangle stationery numbering more than 14,000,000 and its
entertainments, lectures and motion picture shows were attended by
5,000,000 men. It established and carried on thousands of
educational classes, French being the most popular study. Its work
was especially valuable in the education of illiterates and of
foreigners who did not understand English. Some 50,000 who could
not read or write when they entered the training camps received in
this way the rudiments of a common school education. On troop
trains and transports the “Y” workers were present, giving whatever
service the conditions made possible.
Overseas the hut of the Red Triangle was to be found wherever
there were American fighting men—in England, Ireland, Scotland, in
France and Italy, Russia and Siberia, from Gibraltar to Vladivostok,
from the Caucasus to the Murman coast. Sometimes the “hut” was a
dugout, sometimes a ruined chateau, again it was a freight car on a
siding, or a temporary shack, or a substantial building. But, whatever
its form and appearance, it stood for home, for the democratic social
fabric for which the men were fighting, and within it they could
always find light and warmth, cheer and good fellowship, books,
games, music, entertainment, smokes and toothsome dainties.
Motion picture films for the Y. M. C. A. to the average length of
fifteen miles were shipped every week, and at its moving picture
shows there was an average weekly attendance of 2,500,500.
Scores of actors and actresses canceled their engagements and
went overseas to interest and amuse the soldiers and sailors with
performances of all kinds on the hut circuit, organized and directed
by the Over-There Theater League, under the Y. M. C. A. During the
latter months a hundred performances daily, on the average, were
put on in the various camps. None of the players received a salary
and shows of all kinds were free. There were concerts, lectures,
readings, as well as movies and every kind of theatrical
performance. A department of plays and costumes maintained in
Paris sent out to the camps facilities for amateur performances and
fifty professional coaches went from the United States to encourage
and train the soldiers to produce entertainments of their own. Violins,
banjos, mandolins, ukeleles and cornets were sent over by the
thousands, to say nothing of smaller instruments and sheets of
music.
To provide for athletics and physical recreation for the soldiers and
sailors overseas the Y. M. C. A. expended more than a million and a
half dollars. It sent over 1,200 sports leaders and its shipments
included huge quantities of baseballs and bats, boxing gloves,
footballs, ping-pong balls, racquets, nets, tennis balls, running
shoes, and all the paraphernalia of indoor and outdoor sports, to the
value of $2,000,000, which were free for the asking.
The post canteens of the army were taken over by the Y. M. C. A.,
at the urgent request of the commander of the American forces and
against its own desire, and operated throughout the war. This
entailed the running of a huge merchandising proposition foreign to
its customary activities and the work was assumed in addition to its
chosen program of fostering the morale and cherishing the welfare of
the fighting forces. For this post exchange service it furnished
buildings and service without charge and sold to the soldiers at cost
goods to the value of $3,000,000 per month. Its workers often
carried packs of goods into the trenches and distributed them freely.
Because it was all a question of service the organization itself bore
the very considerable loss at which it operated the canteens.
A system of “leave-areas” conducted by the Y. M. C. A. provided
recreation for the men on the seven days’ furlough given to each one
after four months of service. It was not thought desirable by the
military authorities to turn the men loose for their holiday and
therefore several resorts were taken over to furnish interesting
places for them to visit and were put into the hands of the Y. M. C. A.
as hosts and entertainers. Aix-les-Bains was the first and twenty-five
others were added until the men had a wide range of selection
ranging from famed resorts in the Alps to others on the shores of the
Mediterranean. It was a kind of entertainment that had to be created,
for it was entirely without precedent. Largely in the hands of women
workers in the Y. M. C. A., they and their men helpers and advisers
bent their utmost endeavor, resourcefulness and loving care to the
work of giving the men a good time and sending them back to their
duties at the end of their leaves physically and mentally refreshed.
Each area had its athletic field in which every day there were sports
going on and there were mountain climbs, picnics, bicycle rides, and,
in the evening, movies, theatrical entertainments, concerts, music
and dancing.
The women’s contingent of the Y. M. C. A. did effective work both
in these leave areas and in the canteens. Their service was not
enlisted until a year after our entrance into the conflict, but at the end
of hostilities a thousand women were engaged in it, and so insistent
was the call for them that they were recruited as rapidly as possible,
a thousand more being sent over during the next three months. They
were given a week or more of intensive training before sailing to fit
them for the duties they would have to undertake.
Unique in all army as well as in all educational history was the
great educational system which the Y. M. C. A. undertook to
establish, under the authority and with the coöperation of the War
Department. Beginning in the home camps, it was carried across the
sea, developed more and more as time went on, and found its climax
in the “Khaki University.” The final and complete plans were ready
only in time for use with the Army of Occupation in Germany and in
the camps abroad and at home in which the men waited for
demobilization, when $2,000,000 worth of text-books had been
ordered for the work. Some of the foremost educational experts of
the United States, numbering several hundred, were engaged in the
organizing and supervision of the system and many hundreds of
others, members of the alumni and faculties of American educational
institutions who were enrolled among the fighting forces, undertook
the work of instruction. The scheme enabled soldiers and sailors to
continue their studies without expense, whether they desired
elementary, collegiate or professional instruction or agricultural,
technical or commercial training. The scheme, which was finally
taken over by the Army, is described at more length in the chapter on
“The Welfare of the Soldiers.”
So successful and important was the work of the Y. M. C. A. with
the American forces that both the French and the Italian
Governments requested it to establish service centers with their
respective armies. This it did, the American workers who initiated
and supervised the program of recreation and fostering of morale
being assisted, in the respective armies, by French and Italians.
A Pleasant Evening in a Hostess House

Salvation Army Lassies at the Front
The prodigious program of the Y. M. C. A. with the American
forces, which it has not been possible to more than outline, was
carried through largely by volunteer workers who wished to
undertake it as the best way in which they could help to win the war.
Men who were too old to fight or were physically unfit for military
service joyfully welcomed the opportunity to do something that would
aid the fighting men. Many gave up large salaries and left their
situations for the sake of this important service. Others who were
financially unable to leave dependents accepted for them an
allowance much smaller than they could have earned themselves
and gladly took up the work upon the mere payment of their
expenses.
The “Y” workers were on the troop trains that carried the men from
their homes to the training camps and the Red Triangle was at the
fighting man’s side from that moment until he was ready to go over
the top. And sometimes the “Y” worker even went forward in the
charge with the men for whose welfare he was giving his service.
Shell fire not infrequently destroyed the trucks upon which the goods
of the Y. M. C. A. were being carried to the front, its huts were
sometimes shattered in the same way and nine of its workers, two of
them women, were killed by bursting shells. Fifty-seven died in the
service, most of them from wounds, over-work and exposure.
Twenty-three were seriously injured or gassed. Of its workers 152
received official recognition for distinguished services, to thirteen of
whom was awarded the Croix de Guerre and to fifty more other
famous decorations.
The American Army was a reading and thinking army and that one
of the seven great big-brothering organizations which undertook to
supply it with reading matter, the American Library Association, was
kept busy. The Library War Service of the Association had in each of
forty-eight large army and navy training camps and in seventy
hospitals in the United States a central library building, or library
quarters, with branches and stations radiating all over the camp or
hospital area to render its volumes easy of access. It had collections
of books in nearly two hundred hospitals and Red Cross Houses. It
equipped with these collections over five hundred military camps and
posts and aviation fields, schools and repair depots. It supplied with
libraries 260 naval and marine stations and 750 vessels. It had
nearly 2,000 branches and stations placed in Y. M. C. A. and K. of C.
huts, barracks and mess halls. It shipped overseas 2,000,000 books
and 64,000 magazines and distributed 5,000,000 magazines
donated by the public through the mails. In its war service libraries
there were over 5,000,000 volumes. Three hundred and forty trained
librarians supervised its service. Accepted books to the number of
4,000,000 were given by the American people, who provided also
the money with which were bought 1,300,000 more. Book donations
were well sifted before the books were accepted for war service and
the authorities of the association estimated that probably twice as
many were given as were finally used.
But even these enormous quantities of books and magazines were
no more than sufficient to meet the desire for reading shown
throughout the Army and the Navy. The Library War Service of the
Association did its best to supply to every fighting man in the training
camps at home, on the transports, on the cruisers and battleships, in
the stations overseas, in the camps and rest billets, the book he
needed when he wanted it, whether it was light fiction, or a technical
treatise, or a work of history, economics, philosophy or travel. It
supplied books in practically all the modern languages—about forty
were represented in each of the large camps—for both study and
reading and its lists were filled with titles of scientific, technical and
other works that covered the whole range of modern knowledge and
activity, philosophy, literature, history, biography, poetry, art, music,
fiction, drama, economics, sociology, business, travel. There was
demand for them all. Toward the end of the war and after the
armistice the Library War Service bent its energies to meeting the
greatly increased call for vocational books that would enable the
fighting man to become more efficient in his special job or to get a
better one when he should presently be returned to civil life.
To support this vast enterprise of big-brothering the Army the
American people gave without stint to the organizations by which the
work was systematized and carried through. They gave money and
effort and thought and love, because it was for “our boys.” They
responded with more than was asked by each organization in its
separate appeals made during the first year and a half of our war
effort. Then, in order that the appeal for funds might be made more
efficiently and economically, the seven chief organizations united in a
great, nation-wide drive, the money that was subscribed to be
divided proportionately among them. They asked for $170,000,000.
All the preparations had been made for it before the armistice was
signed and it began on that day. Every one believed that the war was
over, but because “our boys” were still overseas and for many weeks
to come would need care, recreation, comforts and entertainment,
no hand withheld its gift. When the week’s drive was over it was
found that $203,179,000 had been subscribed to continue the work
of big-brothering the fighting forces.
CHAPTER XXXII
RUNNING THE RAILROADS
During the first nine months of our participation in the war the
railroads did their best to meet the unusual and mounting demands
upon their facilities and methods. But the entire railroad system had
developed under the principle of competition and, composed as it
was of so many diverse parts and divergent interests, all
accustomed by theory, tradition and practice to competitive methods,
it presently became evident that the coördinated management and
coöperative effort demanded by the emergency would be impossible
under continued private control. The immense increase in traffic
caused by war conditions had strained the existing system to its
utmost effort, and had resulted by the autumn of 1917 in hopeless
congestion of freight at eastern terminals and along the railway lines
far inland. There had been such rapid increase in operating
expenses that the financial situation of the railroads was very bad,
and, under the general financial conditions of the time, had become
a serious menace. The country was at war and its first and most
pressing duty was to prosecute that war to early and complete
victory, which it could not do under the paralysis that was threatening
the transportation system.
For the Government to take control of the railroads was an almost
revolutionary procedure, so opposed was it to American economic
theory, conviction and practice. But the problem was rapidly being
reduced to the bare alternatives of governmental railroad control or
the losing of the war, or, at least, its long-drawn out continuance. But
one solution was possible, and, disregarding all theory and all deeply
rooted custom, the President, in accordance with powers already
conferred upon him by Congress, took possession and assumed
control of the entire railroad system of the United States at the end of
December, 1917.
Management of transportation by rail and water was thereupon put
into the hands of a Director General of Railroads, who thus found
himself at the head of more than 265,000 miles of railway, many
times the mileage of any other nation, and of 2,300,000 employees.
There were about 180 separate operating companies having
operating revenues of $1,000,000 or more per year each and several
hundred more with less than that yearly revenue. The Railroad
Administration, which decentralized its work by dividing the country
into districts, each under a regional director, began its task in the
face of weather conditions without parallel in the history of the
country, which had already almost paralyzed transportation and were
to continue for ten weeks longer.
There was a shortage of freight cars and of locomotives and the
railroads, in common with all the country, were menaced with a
shortage of coal, due mainly to the immensely increased demand
and the breakdown of transportation. So great was the congestion of
freight that in the area north of the Potomac and Ohio Rivers and
east of Chicago and the Mississippi there were 62,000 carloads
waiting to be sent to their destination, while along the lines west and
south of that area there were over 85,000 more carloads held back
by this congestion. Nearly all of it was destined for the eastern
seaboard north of Baltimore.
In addition to the usual transportation business of the country,
hundreds of thousands, mounting into the millions, of soldiers had to
be carried from their homes to cantonments and from cantonments
to ports of debarkation and billions of tons of munitions, food,
supplies and materials of many kinds had to be carried from all parts
of the country upon lines that converged toward eastern ports, while
the immense war building program of the nation—cantonments,
camps, munition plants, shipyards and ships, warehouses, structures
of many sorts—called for the transportation of vast quantities of
material.
By the first of the following May practically all of this congestion
had been cleared up and through the rest of the year there was no
more transportation stringency, although traffic grew constantly
heavier until the end of hostilities. It will illumine the conditions under
which the Railroad Administration achieved its results to mention a
few of its items of transportation. During the ten months ending with
October it handled 740,000 more cars of bituminous coal than had
been loaded during the same period of the previous year. From the
Pacific Northwest there were brought, from April to November, for
the building of airplanes, ships and other governmental activities and
for shipment overseas, 150,000 cars of lumber. During the year
630,000 cars of grain were carried to their destination, the increase
from July to November over the previous year being 135,000 cars.
Livestock movement was especially heavy, showing in all kinds a
large increase. Five hundred and sixty thousand carloads of material
were moved to encampments, shipyards and other Government
projects. From the middle of May to the end of the year the car-
record office showed a total movement of 1,026,000 cars, an
average of 5,700 daily.
Comparison of the physical performance of the roads during the
first ten months of 1918 with that of the similar period in 1917,
reduced to fundamentals, showed an increase in the number of ton-
miles per mile of road per day, in number of tons per loaded car, in
number of tons per freight train mile, in total ton-miles per freight
locomotive per day. The constant purpose was to keep each
locomotive and car employed to its capacity and to make each
produce the maximum of ton-miles with the minimum of train,
locomotive and car miles.
Highly important among the achievements of the Railroad
Administration was the movement of troops. From the first of the
year until November 10th there were transported over the roads
6,496,000 troops, an average of 625,000 per month, the troop
movements requiring 193,000 cars of all types, with an average of
twelve cars to the train. Outstanding features of the troop movement
were that 1,785,000 men were picked up from 4,500 separate points
and moved on schedule to their training camps, that 1,900,000 were
brought into the crowded port terminals for embarkation without
interference with the heavy traffic of other kinds already being
handled there and in the adjacent territory, that 4,038,000 were
carried an average distance of 855 miles, undoubtedly the largest
long distance troop movement ever made. During one period of thirty
days over twenty troop trains were brought each day into the port of
New York. During the entire period from January to November
including these huge troop movements there were but fourteen train
accidents involving death or injury to the men.
To all the necessities of the wartime effort of the railroads—the
enormously increased quantities of freight that had to be moved
expeditiously and the transportation of troops—was added a
considerable increase in the ordinary passenger traffic.
Notwithstanding the earnest and repeated requests of the Railroad
Administration that only necessary journeys should be taken by
civilians, a request that was, indeed, very generally heeded, and the
increase in passenger rates, the passenger traffic all over the
country was much heavier than in any previous year, the increase
amounting in the region east of Chicago to twenty-five per cent.
The efficient handling of all this enormous freight and passenger
traffic was made possible by the policies that were adopted. The
handling of the whole vast network of railroads as one system
eliminated competition and the wasteful use of time, effort and
equipment. The previous usage of the roads in accepting freight at
the convenience of the consignor without regard to the ability of the
consignee to receive it had resulted in the appalling congestion of
terminals and lines in the autumn of 1917. The Railroad
Administration based its policy upon the principle that the consignee
must be considered first and that if he could not receive the freight it
was worse than useless to fill up switches and yards with loaded
cars. In order thus to control traffic at its source a permit system was
adopted which prevented the loading of traffic unless there was
assurance that it could be disposed of at its destination. This policy
proved to be the chief factor in the ability of the transportation
system to meet the enormous demands upon it.
Modification of demurrage rules and regulations induced more
rapid unloading of cars and their quicker return to active use.
Consolidation of terminals, both freight and passenger, greatly
facilitated the handling of cars. Locomotives that could be spared
were transferred from all parts of the country to the congested
eastern region. Coördination of shop work increased the amount of
repairs upon equipment that could be done and kept locomotives
and cars in better condition while new ones were ordered and work
upon them speeded. Rolling stock and motive power were
economized by doing away with circuitous routing of freight and
sending it instead by routes as short and direct as possible, a policy
which saved almost 17,000,000 car miles in the Eastern and
Northeastern Region.
A plan was devised for making up solid trains of live stock and of
perishable freight and also consolidated trains of export freight at
Western points and forwarding them on certain days of the week
directly and rapidly to their destinations. Passenger trains that had
been mainly competitive and such others as could be spared were
dropped, resulting in the elimination during the first seven months of
Federal control of 47,000,000 passenger train miles—an economy in
motive power and equipment without which the successful
movement of troops would have been impossible. Equipment was
standardized, making possible its universal use, and freight cars
were more heavily loaded. In place of the separate ticket offices
made necessary by private and competitive ownership consolidated
ticket offices were opened in all large cities, 101 of these doing the
work of the former 564. The result aimed at was both economy and a
better distribution of the passenger traffic.
The Railroad Administration saw in the inland and coastal
waterways and the coastwise shipping service an important possible
aid in its task of making transportation equal to wartime needs, and
so mid-Western rivers and Eastern canals were brought into
coöperation with railway service and several coast-wise lines of
steamships were made a part of its facilities.
The rental, or return, guaranteed to the railroad companies
amounted for the year approximately to $950,000,000. Upon the
advice of a commission appointed to investigate the matter of wages
and living costs among railroad employees, wages were raised and
threatened labor trouble thereby averted, the increase amounting to
between $600,000,000 and $700,000,000 for the year. In the ten
months ending November 1st the railroad receipts from freight,
passenger and other sources aggregated over $4,000,000,000 and
were almost as large as for the whole of the previous year. The
receipts were greater by 20 per cent, but operating expenses also
had increased by more than $1,000,000,000, the year 1918 breaking
all records for both revenues and expenses. The increase in wages,
in cost of coal, and in all maintenance and operating costs was
responsible for the increase of expenses, which would have been
much greater but for the economies introduced. Freight rates were
raised during the year to help meet the raise of wages, while a
substantial increase in passenger rates was put in force both to help
in that result and to discourage unnecessary passenger traffic during
wartime conditions. There was a final balance against the
Government, as between the net income of the roads and the
guaranteed return to their owners, of between $150,000,000 and
$200,000,000.
The sole purpose of the Government in taking over control of the
railroads was to achieve a more efficient prosecution of the war by
more rapidly forwarding our own war effort and by giving more
effective coöperation to our war associates. Thus, early in the winter
of 1918 the Western Allies made it known to the United States
Government that unless the food promised by the Food
Administration could be delivered to them very soon they could not
continue their war effort. This was immediately after the Railroad
Administration had taken charge of the railroads and was struggling
with the freight congestion extending through the eastern half of the
country, with coal shortage and blizzard weather. Every possible
facility of the Railroad Administration and of the roads it was
operating was brought to the emergency, and railroad officials and
employees worked day and night, with the results that by the middle
of March all the available vessels of the Allies had been filled with
food and dispatched across the Atlantic, while at Eastern seaports
were 6,000 more carloads ready for later shipment.
In carrying out this war-furthering purpose the Railroad
Administration coöperated constantly with the other war
administrative and war prosecuting agencies of the Government, the
Food and Fuel Administrations, the War Trade and War Industries
Boards, the Shipping Board, the Army and Navy Departments. Just
as food, fuel, trade, industry, labor were each and all mobilized for
war effort and all brought into harmonious and effective teamwork,
so the transportation agencies were all bent, first of all, to the same
purpose. Roads, motive power, freight and passenger equipment
were devoted first to the necessities of carrying men from homes to
cantonments and camps and thence to ports of embarkation and of
moving food, munitions, supplies and raw materials to camps, to
shipment points and to places of manufacture for war purposes.
After these war needs were met whatever remained of transportation
facilities was at the disposal of the ordinary commercial traffic of the
country.
In order that the public might better understand the situation and in
order also to better the service of the roads there was instituted a
Bureau of Complaints and Suggestions which dealt with all
dissatisfactions and considered suggested improvements. A very
large number of the railroad employees of all kinds, efficient through
years of service, joined the fighting forces of the nation or engaged
in work more directly concerned with the war and so made it
necessary to fill their places with untrained help. To remedy this
condition training schools were established with successful results.
In the summer of 1918 all express companies were combined and
placed under the management of the Railroad Administration and a
little later telegraph and telephone companies, because of their
refusal to accept an award of the War Labor Board, were unified and
placed under the control of the Postmaster General, as, in the
autumn, was done also with the cable companies.
CHAPTER XXXIII
THE WORK OF WOMEN FOR THE WAR
While the women of the United States did not enter war service by
means of work in industries and auxiliary organizations to the extent
of their enlistment in England, because the man-power problem had
not yet, at the end of hostilities, become serious in this country, the
many and varied kinds of work for the war in which they did engage
was of great importance and it had the devoted and enthusiastic aid
of almost every woman and girl throughout the land. From the
mother who sent her sons across the ocean to the little Girl Scout
who ran errands for a Red Cross chapter, they were ready for any
sacrifice it should be necessary for them to make and any service
they could render. Their spirit was as high, their patriotism as ardent
and their wish to serve as keen as that of their husbands, fathers
and brothers, and their spirit and their service were essential factors
in the war achievements of America. Their spirit was always the
same, but their services were of the greatest variety, being, for the
greater part, such as they could render without leaving their homes.
Being undertaken in addition to their usual duties in the care of
homes and families, their war labors were less outstanding and
much less likely to impress the superficial observer than if they had
been detached from woman’s usual environment. But they were
none the less essential.
The shutting down or curtailment of non-essential industries and
the rapid expansion of those directly or indirectly engaged in war
production shifted many women already possessing some degree of
industrial training into war work plants of one sort or another, while
the need for workers and the desire to give service of direct
consequence led many women to enter factories who had not before
undertaken industrial work. Among the latter class were many of
collegiate education, or of independent means, or engaged in office
work who were moved by patriotism to undertake factory work for the
war. The flow of women into war industrial work increased steadily
throughout the year and a half of our participation and would have
been very greatly augmented if the war had continued long enough
to call the men of the second draft from their situations.
By the end of September, 1918, women were working in munition
plants of many kinds, making shells, grenade belts, fuses, gas
masks, metal parts of rifles, revolvers and machine guns, and many
other sorts of the direct supplies of war. Accurate statistics of their
numbers made in the early summer of 1918 showed that about
1,500,000 women were engaged in the industrial work directly or
indirectly connected with the Government’s war program, while
subsequent estimates added about 500,000 to that number to cover
those entering such work down to the signing of the armistice.
Woman’s Land Army Members Sorting and Grading

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pollutants. The degradation of lignin or hydrocarbons are not mentioned, because

PART II: APPLICATIONS

PART III: USEFUL TOOLS

The Role of the Filamentous

Table 1.1 Filamentous fungi used in bioremediation.

Class Fungi Use References

Geotrichum sp. Azo dyes biotransformation Máximo et al. (2003)

Table 1.1 contd. ...

...Table 1.1 contd.

Class Fungi Use References

Mucor plumbeus Pentachlorophenol degradation Carvalho et al. (2011), (2013)

such as the white-rot fungi species that include Phanerochaete, Bjerkandera,

Enzyme Mechanism of action

Santiestebán et al. 2011, Voběrková et al. 2018). Immobilization of enzyme could be

Cserjesi, A. J. and E. L. Johnson. 1972. Methylation of pentachlorophenol by Trichoderma virgatum.

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