Biotechnology Advances 58 (2022) 107905

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Biotechnology Advances
journal homepage: www.elsevier.com/locate/biotechadv

Research review paper

Antimicrobial properties and applications of metal nanoparticles

biosynthesized by green methods
Olga Maťátková *, Jana Michailidu , Anna Miškovská , Irena Kolouchová , Jan Masák ,
Alena Čejková
University of Chemistry and Technology, Prague, Department of Biotechnology, Technická 5, Prague 6, 166 28, Czechia

A R T I C L E I N F O A B S T R A C T

Keywords: There is a growing interest in the potential and application of metal nanoparticles across many fields. A vast
Metal nanoparticles biosynthesis array of techniques for metal nanoparticle synthesis has been discovered; however, sustainability, cost-
Green methods effectiveness, and environmental concerns favor the green biological approach, using various plant and micro­
Antimicrobial properties
bial sources. This review describes the diversity in green methods for nanoparticle biosynthesis, antimicrobial
Nanoparticles application
Limitations and safety
properties of metal nanoparticles and their potential applications. Metal nanoparticle biosynthesis by extracts
and solutions obtained from plants, bacteria, fungi and templates such as viruses are discussed. As biosynthesized
nanoparticles have been proven to possess antibacterial, antifungal, and even antiviral properties, these are
discussed in detail, with silver and gold nanoparticles as the most studied and with the highest potential for
medical application. The focus on prospective antimicrobial applications of nanoparticles stems from the arising
resistance of many serious pathogens to traditional disinfectants and antibiotics. Other fields for the application
of biosynthesized nanoparticles are also stated briefly, such as in agriculture as pesticides, in wastewater
treatment and bioremediation. Finally, the limitations and safety issues connected with widespread use of
nanoparticles are discussed.

1. Introduction
The field of nanotechnology concerns the engineering, production
and application of materials in the scale range of 1 to 100 nm. During the
last decades, nanotechnology has become one of the most researched
fields due to its broad potential and many applications in various in­
dustrial fields. The number of research publications in the last 20 years,
based on Web of Science search with keywords “biosynthesis” and
“metal nanoparticles” steadily increased during the timeframe, with a
maximum of 612 publications in 2019. This research interest in bio­
logical methods of metal nanoparticles (NPs) synthesis is demonstrated
in Fig. 1.
Nanoparticles often exhibit unique physio-chemical properties,
differing significantly from their macroscale counterparts. These dif­
ferences and subsequent application potential are reflected in their op­
tical, mechanical, chemical, magnetic, and electrical properties (He
et al., 2008; Khan and Fulekar, 2016; Kimber et al., 2018; Valizadeh
et al., 2012), but also in their biological activity, such as antimicrobial or
antiviral (Dikshit et al., 2021; Salem and Fouda, 2021). In the nanoscale,
the surface to volume ratio changes considerably, which results in
higher reactivity in comparison with the parent material (Sahoo et al.,
2021). The material composition of NPs can include various metals (and
metal oxides and salts), non-metal elements (such as carbon nanotubes,
graphenes, and fullerenes), organic nanomaterials (e.g. lipid based NPs)
or hybrids and composites (polymers, ceramics, cements etc.) (Khar­
issova et al., 2019).
Nanoparticles can be formed in natural environment, via abiotic and
biotic mechanisms, including precipitation, erosion, combustion or
volcanic activities (Sharma et al., 2015). Although the natural routes of
NPs synthesis are being explored, many methods were devised to create
synthetic NPs, including physical, chemical and biological approaches,
see Fig. 2 (Maurer-Jones et al., 2013; Salem and Fouda, 2021). These
methods can be classified as either “top-down” or “bottom-up” (Das
et al., 2017; Thakkar et al., 2010).

Abbreviations: DNA, deoxyribonucleic acid; MBC, minimum bactericidal concentration; MFC, minimum fungicidal concentration; MIC, minimum inhibitory
concentration; NPs, nanoparticles; OD, optical density; RNA, ribonucleic acid; ROS, reactive oxygen species.
* Corresponding author.
E-mail address: olga.matatkova@vscht.cz (O. Maťátková).

https://doi.org/10.1016/j.biotechadv.2022.107905
Received 14 September 2021; Received in revised form 15 December 2021; Accepted 7 January 2022
Available online 11 January 2022
0734-9750/© 2022 Elsevier Inc. All rights reserved.
O. Mať átková et al.
The top-down approach usually encompasses nanoparticle produc­
tion by size reduction from bulk material and is the basis of most
physical methods (such as pulsed laser ablation, spray pyrolysis,
evaporation-condensation, ball milling, plasma arc, vapor and gas phase
and lithography (Iravani et al., 2014; Kharissova et al., 2019; Shnoudeh
et al., 2019)). The methods which produce NPs from smaller building
blocks (such as atoms and molecules) belong to the bottom-up approach
and can be found among chemical synthesis methods (e.g. micro­
emulsion, coprecipitation, chemical reduction, electrochemical, micro­
wave, sonochemical, solvothermal and thermal decomposition) and
green biological synthesis methods, which are based on elimination of
hazardous substances via employment of biological catalysts, such as
plant extracts or microbial cell-based systems (see section 2). Biological
synthesis methods using whole organisms or biological molecules bring
significant advantages over physical and chemical methods as they
represent non-toxic and more sustainable approaches to NPs synthesis
(Das et al., 2017; Salem and Fouda, 2021). The green chemistry
approach has been defined as a “design of chemical products and pro­
cesses to reduce or eliminate the use and generation of hazardous sub­
stances”, with its main aim being the modernization of traditional
chemistry processes with a focus on environmental issues, efficiency and
economy (de Souza et al., 2019; Horton, 1999; Salem and Fouda, 2021).
Green synthesis methods are usually regarded as environmentally-
friendly, biocompatible processes, usually conducted at mild pH (4–9)
(Akl et al., 2020; Desai et al., 2020; Kredy, 2018), pressure (atmospheric
pressure) (Vijayaraghavan and Ashokkumar, 2017; Voeikova et al.,
2020), and temperature (room temperature) (Lade and Shanware,
2020), avoiding hazardous materials and conditions and also cost
effective (Salem and Fouda, 2021).
This critical review concerns biological (“green”) methods of metal
NPs synthesis using plant material and extracts, bacteria, fungi, algae
and even viruses and other templates and discussed their properties and
applications. The antibacterial, antifungal and antiviral properties are
discussed in detail and the interconnection of antimicrobial properties
Fig. 1. Web of Science analysis of research publications on the biosynthesis of metal nanoparticles in the last 20 years.
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Biotechnology Advances 58 (2022) 107905
with the NPs source is the main focus of this review. Additionally,
various applications and limitations of NPs are discussed, as well as
perspectives, challenges and ongoing research into as yet unanswered
questions regarding the environmental toxicity of various NPs.
2. Green NPs Synthesis
In NP production, green methods employ substances with a biolog­
ical origin as reducing agents (metal ion reduction), capping agents and
stabilizers (shape, size control and prevention of agglomeration) and as
ligands (coating of NPs surface, metal passivation) (Kharissova et al.,
2019). Biological reducing agents can include small molecules like H2,
but they are more commonly larger water soluble biomolecules such as
reducing polysaccharides, various enzymes, amides, ketones, aldehydes,
carboxylic acids, polyphenols, flavonoids, alkaloids and terpenoids
(Jeevanandam et al., 2016; Nasrollahzadeh and Sajadi, 2015a; Prabhu
and Poulose, 2012). Polysaccharides, proteins and peptides are consid­
ered suitable capping agents and stabilizers, while typical ligands
include amines, thiols, and phosphines (Kharissova et al., 2019). These
biological agents can include complex mixtures or individual com­
pounds (vitamins, saccharides, polyphenols) derived from plants (whole
plants, plant products, extracts) or microorganisms (whole cell, cell-free
extracts; bacteria, fungi and actinomycetes, algae etc.), in both intra­
cellular and extracellular settings in both living and inactivated cells
(Ahmad et al., 2019; Mohamed et al., 2019; Varma, 2012), see Fig. 3.
The use of templates such as biomolecules, DNA, membranes, and vi­
ruses has also been reported (de Souza et al., 2019; Lee et al., 2002;
Rafique et al., 2017). NPs could be synthesized using single bio­
molecules from nature origin, for example various noble metal nano­
particles were synthesized using glutathione (Ag, Au, Pt, Pd) or
riboflavin (Au, Pt) (Baruwati et al., 2009; Nadagouda and Varma, 2006).
The biomolecules in this instance serve as reducing and stabilizing
agents.
The biological synthesis of NPs is carried out by combining a metal
O. Mať átková et al.
precursor compound (salts, acids) with a biological agent under specific
conditions (Nath and Banerjee, 2013). Studies on biological synthesis of
NPs using various biological agents include metals and their derivative
compounds, often aiming to produce Ag, Au, Zn, Ni, Pd, Pt, Se, and Cu
nanoparticles (Dikshit et al., 2021). Specifically, metal compounds such
as AgNO3, HAuCl4, (CH3COO)2Zn, (NiNO3)2FeCl3, PdCl2, H2PtCl6,
Na2SeO3 and Cu(NO3)2 are commonly used for the synthesis of their
respective metal NPs (Dikshit et al., 2021). The presence of specific
suitable biomolecules results in the reduction, capping, and stabilizing
of the parent metal compound in the form of NPs (Dikshit et al., 2021;
Kuppusamy et al., 2016).
The technological implementation of the biological NPs synthesis
depends mainly on the type of bioagent. For example, the synthesis of
metal NPs using plant extracts usually begins with collecting and
washing the desired part of the plant (leaf, fruit, peel, etc.) with distilled
water, followed by drying, crushing and finally extraction into water/
solvent (most often ethanol or methanol) (Mohamad et al., 2014). The
solid plant residues are then separated from the extract by filtration or
centrifugation and mixed with the metal precursor compound under
certain conditions. The reaction time (and thus the characteristics of
nanoparticles) is influenced by the ratio of reactant concentrations,
temperature, pH, light, ultrasound or microwave heating, which are
among the leading parameters modified to achieve optimal NPs syn­
thesis (Mittal et al., 2013; Varma, 2012). After incubation, the nano­
particles are usually centrifuged at high speed, washed thoroughly in
water/solvent (ethanol, methanol), and collected (Singh et al., 2016),
alternatively they can be applied without separating the extract. The
amount and morphology (shape, size distribution) of the resulting
nanoparticles are greatly influenced by the reaction conditions. The
source from which the extract is prepared has also a major influence,
because individual plant extracts can differ significantly in content and
combination of reducing and stabilizing biomolecules (Kumar and
Yadav, 2009).
It is important to address quality of research in the context of green
methods as there definitely seems to be room for improvement. There
are many studies where insufficient data about the production method is
given e.g., specification of reagent ratios, reaction conditions etc. In­
formation about sample preparation for characterization is not present
in most studies. Moreover, results from analyses are often poorly com­
mented and are not discussed sufficiently in several present studies.
Finally, determination of concentration is a severe issue in studies
concerning biosynthesis of metal nanoparticles, with some studies tak­
ing scientifically unsound shortcuts, such as calculating nanoparticle
concentration directly from the initial concentration of precursor, thus
assuming 100 % yield. Another issue not often discussed is the
Fig. 2. Routes of nanoparticles synthesis – biological, chemical, and physical methods.
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Biotechnology Advances 58 (2022) 107905
applicability of original research on NPs biosynthesis into their large
scale industrial production, which can be benefited by the use of key
variables determination, e.g. with modelling, such as proposed by Cin­
elli et al. (2015) – see chapter 4.5.1 for further discussion.
The two main biological systems used for the synthesis of metal NPs
are summarized in the following sections – plant and microbial NPs
synthesis.
2.1. Plant-mediated NPs biosynthesis
The biosynthesis of NPs using plants originates in the ability of plants
to avoid toxic effects of excess metals by various mechanisms. Com­
pounds with antioxidant properties play an important role in these
protective processes, as many metals generate ROS whose activity can
result in serious damage to the cell (Kharissova et al., 2019). For
example, plant antioxidant mechanisms can be facilitated by phenolic
compounds. Accumulation of metals and their reduction and stabiliza­
tion inside plant tissues has been studied with respect to bioremediation
applications (phytoremediation). The research into these processes led
to the observation of metal deposits in plants in the form of NPs (Gardea-
Torresdey et al., 2002; Makarov et al., 2014).
NPs synthetized with the use of plant material will differ in their
properties, size and shapes depending on several factors which can be
immutable (presence of compounds in specific plants acting as reducing
or capping agents) and variable (extraction method, synthesis condi­
tions, such as temperature, pH, reaction time, concentration of metal
parent compound etc.) (El-Seedi et al., 2019; Kharissova et al., 2019;
Marslin et al., 2018; Mohammadinejad et al., 2016).
There is a large number of studies reporting on the synthesis of
various metal NPs using many different plants (Das et al., 2017; Dikshit
et al., 2021; Mittal et al., 2013; Shankar et al., 2003; Sheny et al., 2013),
see Table 1. The methods of NPs biosynthesis mediated by plants use
three main approaches; the use of whole plants, plant extracts, and in­
dividual isolated phytochemicals (Dikshit et al., 2021).
2.1.1. Whole plant mediated NPs biosynthesis
The use of whole plants for metal NPs synthesis is based on the up­
take of metals from the environment by the plant and subsequent
intracellular reduction and accumulation in the form of NPs.
This process is assisted by many compounds, such as amino acids,
alkaloids, flavones, proteins, polyphenols, polysaccharides, tannins,
terpenoids, and vitamins (Ahmad et al., 2010; El-Seedi et al., 2019). This
method leads to a great variation in the NP properties due to the
complexity of the process and lower possibility of its regulation. In a
whole plant synthesis, the NPs can be located in various plant parts and
O. Mať átková et al.
Fig. 3. Plant and microbial processes employed for green synthesis of NPs.
cell types; studies reported widespread localization (epidermis, cortex,
and vascular tissue) with the majority of NPs being found in xylem pa­
renchyma cells (Corredor et al., 2009; Marchiol et al., 2014; Sharma
et al., 2007). The localization of NPs significantly influences the
complexity of the subsequent process of NPs isolation and purification.
The use of plants for the preparation of metal NPs had been motivated by
their well-known ability to phytoremediate heavy metals from the
environment. In order to synthesize nanoparticles using whole plants, it
is necessary to enrich the soil or surrounding medium with a soluble
form of the respective metal ions (Iravani, 2011). Whole plant synthesis
of silver nanoparticles was successfully achieved with the use of Mesa
variety alfalfa sprouts grown on silver-rich solid medium (from 40 to
320 mg/L of silver). After 9 days, the sprouts were harvested, freeze-
dried and pulverized into homogenous powder. Characterization of
the sprouts revealed successful production of silver nanoparticles
ranging from 2 to 20 nm in diameter (Gardea-Torresdey et al., 2003).
Synthesis of both silver and gold NPs by Medicago sativa and Brassica
juncea cultivated using aqueous substrate containing AgNO3 (from 1000
to 10 000 mg/L) and KAuCl4 (1000 mg/L), respectively, was reported by
Bali and Harris (2010); Harris and Bali (2008).
2.1.2. Plant extract mediated NPs biosynthesis
The use of plant extracts or individual plant substances utilizes a
simpler approach (in contrast to whole plant biosynthesis) with a po­
tential for higher control and versatility, higher yield, simpler isolation
and scale-up, therefore earning most of the scientific focus in the field of
plant-based NPs biosynthesis (Dikshit et al., 2021; Mittal et al., 2013;
Thakkar et al., 2010).
Extracts of various plants and different parts of said plants have been
used in the synthesis of metal NPs. The plant extracts can be made from
roots, stems, bark, leaves, flowers, fruits, seeds etc. (Dikshit et al., 2021).
Successful biosynthesis of noble metal nanoparticles (Ag, Au, Pt, Pd) was
reported using fruit extracts of blackberry, blueberry and pomegranate,
root extracts of turmeric or beet and aqueous extracts from green tea
(leaves) and coffee (Kou and Varma, 2012a; Nadagouda et al., 2014;
Nadagouda and Varma, 2008). As the management of food and agri­
culture waste products has emerged as a major concern in recent years,
the NPs synthesis using derivatives of agricultural plants rich in bio­
logically active substances present an opportunity for utilization and
exploitation of such materials. Therefore, the plant NPs synthesis can
utilize materials that originate as food or agriculture waste, such as
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Biotechnology Advances 58 (2022) 107905
citrus or banana peel extracts (Bankar et al., 2010; Castro et al., 2015),
aiding the profitability of the life cycle analysis of the total production
cost and conveniently exploiting waste materials as resources. For
example aqueous extract from grape pomace was used for successful
biosynthesis of silver, gold, platinum and palladium NPs (Baruwati and
Varma, 2009).
The plant extracts can be obtained by various methods, such as hot/
cold extraction, Soxhlet extraction and with the aid of different solvents
(Dikshit et al., 2021). The molecular composition of the extract is key for
the viability of the process of metal reduction and NPs capping and
stabilization. However, due to the high complexity and variability of
plant extracts, the respective roles of individual compounds are largely
unknown (Dikshit et al., 2021). The compounds participating in the
reduction of metal ions are known to include hydroxyl, carbonyl, amino,
and methoxide functional groups, all of which exhibit electrostatic in­
teractions with metal ions followed by the reduction reaction (El-Seedi
et al., 2019; Küünal et al., 2018). Important role in the process is played
by plant secondary metabolites, alkaloids, cofactors, enzymes, flavo­
noids, polyphenols, etc. (El-Seedi et al., 2019).
The NPs synthesis using plant extracts is carried out by combining
the plant extract with a solution containing metal ions (utilizing metal
salts or acids) under controlled conditions, such as pH, temperature,
reaction time, oxygen and light exposure, etc. (Dikshit et al., 2021). The
composition of the plant extract and its concentration (as well as the
metal compound concentration) mainly determine the NP morphology,
while the reaction conditions (temperature, pH) are reported to modify
the size of the NPs (Castro et al., 2011; Salayová et al., 2021; Sankar
et al., 2014). For example, spherical AuNPs (2–10 nm) were synthesized
using Nerium oleander leaf extract from dried, ground N. oleander leaves
which were mixed with water (0.1 g/ml) and stirred at 60 ◦ C for 5 hours.
Filtered extract was mixed with chloroauric acid to produce the NPs
(Tahir et al., 2015).
Using Gnidia glauca flower extract, variable shapes (triangular,
hexagonal, spherical) of gold nanoparticles of sizes 50–150 nm were
obtained (Ghosh et al., 2012).
The plant extract synthesis of metal NPs can be divided into several
successive stages. The process starts with the activation step, i.e.
reduction of metal ions to metal atoms, and is followed by the nucleation
of the reduced metal atoms into smallest NPs and further into larger
sized NPs, accompanied by an increase in thermodynamic stability of the
dispersion. It was found that higher temperatures yield larger
O. Mať átková et al. Biotechnology Advances 58 (2022) 107905

Table 1
Plant-synthesized metal NPs and their properties.
NPs Plant species Size (nm) Applications Ref.

Whole plant
Au Brassica juncea 5–10 – (Bali and Harris, 2010)
Medicago sativa 10–20 – (Bali and Harris, 2010)
Plant extracts
Allophylus cobbe leaves 2–10 Antibacterial and antibiofilm (Gurunathan et al., 2014)
Annona reticulata leaves 6.48–8.13 Antibacterial, larvicidal (Parthiban et al., 2019)
Calendula officinalis leaves 30–50 – (Alanazi and Hesham, 2016)
Calendula officinalis seeds 5–10 – (Baghizadeh et al., 2015)
Camellia sinensis 34.68 ± 4.95 (DLS) Antibacterial and cytotoxicity evaluation (Rolim et al., 2019)
Eucalyptus globulus leaves 8–35 Scolicidal activity (Salih et al., 2020)
Hibiscus rosa sinensis leaves – – (Philip, 2010)
Lansium domesticum fruit peel 10–30 Antimicrobial, biocompatibility (Shankar et al., 2014)
Origanum majorana leaves 25–50 – (Zahran et al., 2018)
Ag
Panax ginseng leaves 5–15 Antimicrobial (Singh et al., 2016)
Persea americana seeds 20–40 Antimicrobial (Girón-Vázquez et al., 2019)
Phyllanthus amarus (parts of whole plant) 24 ± 8 Antibacterial (Singh et al., 2014)
Phyllanthus emblica fruit 60–80 – (Dhar et al., 2021)
Piper nigrum leaves 8–35 Scolicidal activity (Salih et al., 2020)
Tabebuia berteroi leaves 2 (average) Catalytic (Vellaichamy and Periakaruppan, 2016)
Ziziphora tenuior leaves 8–40 – (Sadeghi et al., 2015)
Ziziphus Jujuba leaves 20–30 Antibacterial, catalytic (Gavade et al., 2015)
Ziziphus Spina-Christi leaves 8–35 Scolicidal activity (Salih et al., 2020)
Camellia sinensis leaves ~10 Antibacterial (Onitsuka et al., 2019)
Elettaria cardamomum seeds 15.2 (average) Antioxidant, antibacterial, anticancer (Rajan et al., 2017)
Hibiscus rosa sinensis leaves – – (Philip, 2010)
Hygrophila spinosa (parts of whole plant) – Antioxidant and anticancer (Satpathy et al., 2020)
Lansium domesticum fruit peel 20–40 Antimicrobial, biocompatibility (Sankar et al., 2014)
Au
Nigella arvensis leaves 3–37 Antibacterial, antioxidant, cytotoxic, catalytic (Chahardoli et al., 2018)
Panax ginseng leaves 10–20 Antimicrobial (Singh et al., 2016)
Stevia rebaudiana leaves 5–20 – (Sadeghi et al., 2015)
Terminalia chebula seed 6–60 Antibacterial (Kumar et al., 2012)
Zingiber officinale root 5–15 Blood compatibility (Kumar et al., 2011)
Crotalaria candicans leaves 30 (average) Antibiofilm (Lotha et al., 2019)
Ginkgo biloba L. leaves 15–20 Catalytic (Nasrollahzadeh and Sajadi, 2015a)
Cu
Syzygium aromaticum buds 15 (average) Antimicrobial (Rajesh et al., 2018)
Ziziphus spina-christi (L.) fruit 5-20 Antibacterial, removal of crystal violet dye (Khani et al., 2018)
Citrofortunella microcarpa leaves 54–68 Photodegradation of Rhodamine B dye (Rafique et al., 2020)
CuO
Cymbopogon citratus leaves 11.4–14.5 Antibacterial and antibiofilm (Cherian et al., 2020)
Moringa oleifera leaves 3.4–7.4 Antibacterial, removal of nitrate from water (Katata-Seru et al., 2018)
Fe Moringa oleifera seeds 2.6–6.2 Antibacterial, removal of nitrate from water (Katata-Seru et al., 2018)
Trigonella foenum–graecum seeds 7–14 Antibacterial, photocatalytic dye degradation (Radini et al., 2018)
Cinnamomum camphora leaves 3.2–6.0 – (Yang et al., 2010)
Couroupita guianensis Aubl. fruit 5–15 Antibacterial and cytotoxic (Gnanasekar et al., 2018)
Pd Filicium decipiens leaves 2–22 Antibacterial (Sharmila et al., 2017)
Rosmarinus officinalis leaves 15–90 Antimicrobial and catalytic (Rabiee et al., 2020)
Sapium sebiferum leaves – Antibacterial, photocatalytic (Tahir et al., 2016)
Pd/CuO Theobroma cacao L. seeds 40 (average) Catalytic (Nasrollahzadeh et al., 2015)
Ocimum sanctum leaves 23 (average) Water electrolysis (Soundarrajan et al., 2012)
Pt Taraxacum laevigatum (parts of whole plant) 2–7 Antibacterial (Tahir et al., 2017)
Xanthium strumarium leaves 22 (average) Antimicrobial, cytotoxic (Kumar et al., 2019)
TiO2 Euphorbia heteradena Jaub root 17–45 – (Nasrollahzadeh and Sajadi, 2015a)
ZnO Spinacia oleracea leaves 40.59 (average) Effect on Vigna radiata (Lakshmi et al., 2017)

nanoparticles under Ag+ precursor abundance, however, when insuffi­
cient concentration of Ag+ are present in the solution, the trend is not
rising linearly. Conversely, under high temperatures (80-90 ◦ C), smaller
nanoparticles are produced with rising temperature (Liu et al., 2020). As
tempering a solution evenly using heat transfer is often a problem,
several studies use microwaves as a method to provide increased reac­
tion kinetics homogeneously (Baruwati et al., 2009; Hebbalalu et al.,
2013; Kou and Varma, 2012b; Varma, 2012). Another parameter which
can influence the reaction kinetics is UV light. In a study by Filip et al.
(2019), a 6 V UV lamp (365 nm) was used for the photochemical pro­
duction of silver and gold nanoparticles mediated by Cornus mas fruit
extract. Free radicals in the extract were formed by the UV-light and
induced the reduction of Ag+ / AuCl4- ions. Manjamadha and Muthu­
kumar (2016) reported that ultra-sonication can shorten reaction time.
Ultrasound reduces reaction rate and reduction time by increasing the
interactions of functional groups from the biological agent and metal
ions. Lastly, the finishing parts of the process (e.g. capping) determine
the final shape and size of the NPs (Dikshit et al., 2021; Makarov et al.,
2014; Si and Mandal, 2007).
The resulting NPs can be used and stored within the solution
including the remaining plant extract or can be further separated and
purified before application (washing, centrifugation, column chroma­
tography) (Nasrollahzadeh and Sajadi, 2015a).
Various approaches were already successfully employed in the
context of plant extract synthesis. In some studies, whole plant extracts
were used for the production of nanoparticles. Mahitha et al. (2011)
reported the successful synthesis of silver nanoparticles using whole
plant ethanolic extract made from Bacopa monniera. Nanoparticles were
prepared by mixing the extract with a 25 mM aqueous solution of silver
nitrate (1:30) and incubating at room temperature for one hour. This
procedure led to the formation of mostly spherical nanoparticles with a
size of 5–30 nm. Similar approach for plant extract NPs synthesis was
adopted by Vijay Kumar et al. (2014), who reported successful synthesis
of silver nanoparticles with the use whole plant extract made from
Boerhaavia diffusa. Silver nanoparticles were also synthesized using
whole plant extract made from Bacopa monniera (Mahitha et al., 2011)

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O. Mať átková et al.
and Clitoria ternatea (Malabadi et al., 2012). Moreover, biosynthesis of
AuNPs using aqueous extract of Hygrophila spinosa was reported by
Satpathy et al. (2020).
The most frequently reported method utilizing plant extracts seems
to be an approach using leaf extracts. Rabiee et al. (2020) reported the
successful synthesis of palladium nanoparticles using aqueous extract of
Rosmarinus officinalis. The extract was prepared by washing R. officinalis
leaves with distilled water, then drying at room temperature, followed
by cutting into small pieces and grounding. The powder was mixed with
deionized water (10 g/100 ml) and boiled for 15 minutes. Finally, the
extract was filtered to remove all plant residues. Nanoparticle synthesis
was conducted by adding 10 ml of the extract to 20 ml of the palladium
precursor salt and stirring at room temperature for 24 hours. Palladium
NPs in the size range from 15 to 90 nm were obtained. Salih et al. (2020)
reported AgNPs synthesis with the use of leaf extracts from Piper nigrum,
Ziziphus Spina—Christi and Eucalyptus globulus. Dhar et al. (2021) re­
ported AgNPs synthesis by using extract from Phyllanthus emblica. AuNPs
and AgNPs were synthesized using leaf extracts of Crassocephalum rubens
(Adewale et al., 2020). AuNPs, AgNPs, and PdNPs were synthesized
using leaf extract of Rosmarinus officinalis (Rabiee et al., 2020). FeNPs
and PdNPs were synthetized by the leaf extract of Tabebuia berteroi
(Vellaichamy and Periakaruppan, 2016) and Withania coagulans
together with graphene oxide to form a nanocomposite (Atarod et al.,
2016).
As mentioned above, other parts of plants used for the synthesis of
metal nanoparticles include stems, roots, fruits seeds and other plant
parts, preferable from agricultural waste products. In this regard, AgNPs
were synthesized using fruit extract made from Phyllantus emblica
(Renuka et al., 2020) and using stem extract from Phyllantus pinnatus
(Balachandar et al., 2019). Pd/CuONPs were biosynthesized using seed
extracts from Theobroma cacao, rich in phenolic antioxidants (Nasrol­
lahzadeh et al., 2015). TiO2NPs were synthetized by root extract from
Euphorbia heteradena which exhibited the presence of phenolic com­
pounds, used as both reducing and capping agents (Nasrollahzadeh and
Sajadi, 2015b).
2.2. Microbial mediated NPs biosynthesis
Various unicellular systems have been investigated for their poten­
tial NPs production. Reports describe successful exploitation of bacteria,
yeast, actinomycetes, fungi, and algae; metal NPs under study include
Ag, Au, Zn, Ti, Te, Se, Cu, Mg, their oxides, alloys, quantum dots and
other nanocomposites (Fouda et al., 2018; Hassan et al., 2018;
Mohamed et al., 2019; Salem and Fouda, 2021). The disadvantages of
microbial synthesis of NPs include the need for special equipment for
microbial cultivation and its control and maintenance, variability of the
microbial product depending on cultivation conditions and, lastly,
biosafety issues that might arise with some microbial strains (Kharissova
et al., 2019).
Promising candidates for microbial mediated NPs biosynthesis can
be constructed using genetic engineering. Prospective properties of the
genetically engineered organisms can include enhanced resistance to
metal toxicity, high throughput, energy efficiency and ecological pro­
cedures (Iravani and Varma, 2019). Among reported studies are re­
combinant E. coli coexpressing metallothioneins and phytochelatin
synthase (Choi et al., 2018) or heterologous host construction for
biosynthesis of magnetic NPs by transferring genes from Magneto­
spirillum gryphiswaldense to Rhodospirillum rubrum (Kolinko et al., 2014).
There are studies trying to elucidate the mechanisms and compounds
involved in the microbial-mediated NPs synthesis although, as with the
plant extracts, we can expect a high degree of specificity in each mi­
crobial system, depending on the type of microorganism and the con­
ditions of its cultivation (temperature, pH, substrate, etc.) (Narayanan
and Sakthivel, 2010; Salem and Fouda, 2021). The participating com­
pounds include cell wall polymers, quinones, enzymes (such as NADPH-
dependent nitrate reductase), peptides and polysaccharides (as capping
6
Biotechnology Advances 58 (2022) 107905
agents) (Zhao et al., 2018).
The microbial-mediated NP synthesis can utilize both extracellular
and intracellular approaches, using whole cells or cell-free supernatants,
cell lysates, or extracts (de Souza et al., 2019). In comparison with plant
mediated NPs synthesis, the microbial systems include a cultivation step
alternatively amended by a purification step aimed at obtaining the
active agents, which can be derived from various stages of microbial
metabolism – specific examples are stated in following sections.
2.2.1. Bacteria mediated NPs biosynthesis
Bacterial production of NPs is facilitated by various extracellular and
intracellular processes and compounds both during the active growth
phase or using secondary metabolism by-products (de Souza et al., 2019;
Salem and Fouda, 2021). The cell-metal interactions are often based on
the ability of bacteria to suppress their toxic and adverse effects and are
an intrinsic part of the microbial resistance mechanisms to heavy, toxic
metals (Vigneshwaran et al., 2007) e.g. by reducing and precipitating
the toxic metal from the soluble form into insoluble non-toxic metal
nanoparticles (Narayanan and Sakthivel, 2010). The biosynthesis of
metal NPs can be a part of the biomineralization process, using the cell
as a scaffolding, or can take place during active metabolic processes,
such as biosorption, complexation and precipitation or intracellular
bioaccumulation (Iqtedar et al., 2019; Narayanan and Sakthivel, 2010).
The cultivation of bacteria, in contrast to other microbes, is character­
ized by high growth rates and yield, and low requirements for cultiva­
tion conditions. The bacterial biosynthesis of NPs is therefore a flexible
technique with a potential for large-scale production (Salem and Fouda,
2021). Although extracellular production has obvious advantages in
easier down-stream processing, intracellular production has been re­
ported to lead to NPs with higher monodispersity (Bao et al., 2003;
Narayanan and Sakthivel, 2010). Microbial NPs synthesis is conducted
by combining either cell biomass or cell-free extract/supernatant ob­
tained after pre-cultivation and the appropriate metal parent compound
under suitable reaction conditions (time, pH, temperature and pressure)
(de Souza et al., 2019).
Production of AgNPs by bacteria has been one of the first reported
studies on the possibilities of microbial-mediated NP biosynthesis
(Prabhu and Poulose, 2012) and although the focus has been on the
AgNPs, there are reports on the synthesis of many other metal NPs using
a variety of microorganisms (see Table 2). Synthesis of AgNPs by cell-
free extracts (e.g. from Bacillus brevis (Saravanan et al., 2018a), Strep­
tomyces griseoplanus (Vijayabharathi et al., 2018)) and intracellular (e.g.
Lactobacillus fermentum (Sintubin et al., 2009), Pseudomonas sp. (John
et al., 2020)) has been reported. In extracellular synthesis, the first step
is usually cultivation of microorganisms in optimal growth conditions
(temperature, pH, growth medium and others) for 1-2 days. Then the
biomass is separated from the media (by centrifugation or filtration). To
synthesize NPs, the obtained supernatant/filtrate is mixed with the
metal precursor solution. Another option is the use of aqueous cell-free
extract. In this case, the cells obtained as described above are washed
and incubated in sterile water. The biomass is then separated from the
resulting suspension and formed cell-free aqueous extract is mixed with
a solution of metal ions (Kato and Suzuki, 2020; Singh et al., 2016). Wei
et al. (2012) used aqueous extract of Bacillus amyloliquefaciens obtained
by ultrasonication of washed pre-cultured cells and synthesized AgNPs
of sizes 4–24 nm.
Intracellular biosynthesis is done by incubation the cell suspension
with metal precursor for a few minutes to days. To minimize the effect of
the culture medium components, the precultured biomass could be
harvested, washed with sterile water, and then incubated with a metal
salt solution. The biomass of Lactobacillus casei subsp. casei was used as
biocatalyst for the AgNPs (25–50 nm) formation. The reaction mixture
contained harvested biomass, silver nitrate, glucose (as electron donor)
and phosphate buffer (Korbekandi et al., 2012).
Husseiny et al. (2007) synthetized AuNPs using cell-free supernatant
from Pseudomonas aeruginosa. They mixed supernatant after culturing
O. Mať átková et al. Biotechnology Advances 58 (2022) 107905

Table 2
Microbially synthesized metal NPs and their properties.
NPs Species Source Size (nm) Applications Ref.

Bacteria
Bacillus brevis cell-free extract 41–68 Antibacterial (Saravanan et al., 2018a)
Bacillus licheniformis cell-free supernatant ~ 50 nm – (Kalishwaralal et al., 2008)
Bacillus methylotrophicus cell-free supernatant 10–30 Antimicrobial (Wang et al., 2016)
Bacillus sp. biomass 5–15 – (Pugazhenthiran et al., 2009)
Lactobacillus fermentum biomass 5–80 – (Sintubin et al., 2009)
Ag Pseudomonas sp. biomass 20–70 Antimicrobial (John et al., 2020)
Pseudomonas stutzeri biomass 200 – (Klaus et al., 1999)
Rhodococcus spp. biomass 10–12 Antimicrobial (Otari et al., 2015)
Shewanella oneidensis biomass 2–11 Antibacterial (Suresh et al., 2010)
Sinomonas mesophila cell-free supernatant 4–50 Antimicrobial (Manikprabhu et al., 2016)
Streptomyces griseoplanus cell-free extract 19–20 Antifungal (Vijayabharathi et al., 2018)
Bacillus subtilis - 20–25 Catalytic (Srinath et al., 2018)
Escherichia coli DH5α biomass 25 Direct electrochemistry of hemoglobin (Du et al., 2007)
Mycobacterium sp. cell-free supernatant 5–55 – (Camas et al., 2018)
Au Pseudomonas aeruginosa cultural supernatant 15–30 – (Husseiny et al., 2007)
Rhodopseudomonas
cell-free extract 10–20 – (He et al., 2008)
capsulata
Streptomyces sp. cell-free supernatant 90 (average) Antifungal (Gopal et al., 2013)
Kocuria flava cell-free supernatant 5–30 – (Kaur et al., 2015)
Cu
Shewanella oneidensis biomass 20–40 Catalytic (Kimber et al., 2018)
Antimicrobial, antioxidant,larvicidal and
CuO Streptomyces sp. biomass filtrate 78–80 (average) (Hassan et al., 2019)
biocontrol
Pd Shewanella oneidensis - 10–100 Catalytic (Wang et al., 2019)
Jeotgalicoccus coquinae 5.74
cell lysate supernatant Antioxidant and antimicrobial (Eramabadi et al., 2020)
ZC15 (average)
Pt
electrochemically active
Shewanella loihica 1–10 Catalytic (Ahmed et al., 2018)
biofilm
Escherichia coli biomass 2–5 – (Sweeney et al., 2004)
CdS Extracellular polymeric
Pseudomonas aeruginosa 20–40 – (Raj et al., 2016)
substances
Bacillus amyloliquefaciens biomass in culture medium 22.11–97.28 Photocatalytic degradation of dye (Khan and Fulekar, 2016)
Órdenes-Aenishanslins et al.,
TiO2 Bacillus mycoides 40–60
2014
Lactobacillus sp. biomass 8–35 – (Jha et al., 2009)
Bacillus megaterium cell-free extract 45–95 Antimicrobial (Saravanan et al., 2018b)
ZnO
Streptomyces sp. cell-free supernatant 20–50 Antimicrobial and anticancer (Balraj et al., 2017)
Fungi and yeasts
Aspergillus fumigatus cell-free filtrate 5–25 – (Bhainsa and D’Souza, 2006)
Aspergillus terreus cell-free supernatant 1–20 Antimicrobial (Li et al., 2012)
Cryptococcus laurentii cell-free supernatant 35–400 Antibacterial (Fernández et al., 2016)
Fusarium oxysporum biomass 5–13 Antibacterial and cytotoxic (Husseiny et al., 2015)
Ag
Phomopsis liquidambaris cell-free filtrate 18.7 (average) Antimicrobial and larvicidal (Seetharaman et al., 2018)
Rhizopus stolonifer mycelial extract 2.86 ± 0.3 nm – (AbdelRahim et al., 2017)
Rhodotorula glutinis cell-free supernatant 15–220 Antifungal (Fernández et al., 2016)
Trichoderma viridae cell-free filtrate 5–40 Antimicrobial (Fayaz et al., 2010)
Cladosporium
cell-free extract 60 nm Antimicrobial and antioxidant (Joshi et al., 2017)
cladosporioides
Fusarium oxysporum biomass 20–40 – (Mukherjee et al., 2002)
Pichia jadinii biomass < 100 nm – (Gericke and Pinches, 2006)
Rhizopus oryzae cell-free extract 16–43 – (Kitching et al., 2016)
Au
Saccharomyces cerevisiae biomass > 100 nm – (Lin et al., 2005)
Trichoderma harzianum biomass 26–34 Antibacterial (Tripathi et al., 2018)
Trichothecium sp. biomass – – (Ahmad et al., 2005)
Verticillium sp. biomass 12–28 – (Mukherjee et al., 2001)
Yarrowia lipolytica biomass 15 – (Agnihotri et al., 2009)
32 (Sriramulu and Sumathi,
Pd Saccharomyces cerevisiae cell-free extract Catalytic
(average) 2018)
Fusarium oxysporum biomass 10–100 – (Riddin et al., 2006)
Pt
Penicillium chrysogenum cell-free filtrate 5–40 Antibacterial and cytotoxic (Subramaniyan et al., 2018)
CdS Candida glabrata - 20 Å – (Dameron et al., 1989)
Aspergillus niger cell-free filtrate 53–69 Antimicrobial and dye degradation (Kalpana et al., 2018)
ZnO Aspergillus terreus cell-free filtrate 10–45 Antibacterial and UV- protection (Fouda et al., 2017)
Pichia kudriavzevii cell-free filtrate ~ 10–61 Antimicrobial and antioxidant (Moghaddam et al., 2017)
Algae
Polysiphonia cell-free extract 5–25 Anticancer (Moshfegh et al., 2019)
Ag
Sargassum muticum cell-free extract 43–79 Antibacterial (Madhiyazhagan et al., 2015)
Dunaliella salina cell-free extract 1–30 Anticancer (Singh et al., 2019)
Fucus vesiculosus biomass – – (Mata et al., 2009)
Chlorella vulgaris biomass 40–60 – (Luangpipat et al., 2011)
Au
Rhizoclonium fontinale biomass – – (Parial and Pal, 2015)
Sargassum wightii biomass 8–12 – (Singaravelu et al., 2007)
Spirulina platensis protein extract ~ 5 nm Antibacterial (Suganya et al., 2015)
Pd Chlorella vulgaris cell-free extract 70 Catalytic (Mishra et al., 2020)
(continued on next page)

7
O. Mať átková et al.
Table 2 (continued )
NPs Species Source Size (nm)
CuO Bifurcaria bifurcata cell-free extract 5–45
ZnO Chlamydomonas cell-free extract 55–80
reinhardtii
Chlorella cell-free extract 20 ± 2.2
Chlorella vulgaris cell-free extract 20–50
Viruses
bacteriophage - 20–50
Au
tobacco mosaic virus - 10–40
TiO2 20–40
M13 virus - –
(nanowire) (diameter)
the microorganism with chloroauric acid (to give a 1mM solution) and
obtained spherical AuNPs in the range of 15–30 nm. In another study,
Escherichia coli biomass was used to mediate the AuNPs biosynthesis (Du
et al., 2007). The authors mixed washed biomass resuspended in sterile
water with an aqueous solution of chloroauric acid (to give a 1mM so­
lution) and incubate the mixture in laboratory ambient conditions for
120 h. This led to the formation of spherical AuNPs with an average size
25 nm.
Other metal nanoparticles including CuNPs and CuONPs were pre­
pared using bacteria. Kaur et al. (2015) reported the successful
biosynthesis of CuNPs using the supernatant after cultivation of the
marine bacteria Kocuria flava. In this study, the cell-free supernatant was
mixed with copper nitrate and the mixture was incubated at 30 ◦ C for 24
h. The experiment resulted in the formation of spherical and quasi-
spherical CuNPs with an average size ranging between 5 and 30 nm.
Endophytic Streptomyces sp. was used for the biosynthesis of spherical
CuONPs with average size 80 nm by mixing the aqueous biomass filtrate
with CuSO4.5H2O and incubating the mixture at 35 ◦ C for 6 h in the dark
(Hassan et al., 2019).
2.2.2. Fungi mediated NPs biosynthesis
Beside prokaryotic microorganisms, some species of fungi and yeast
have been studied as potential NPs producers. Although their growth
rates are lower than those found in bacteria, they possess some advan­
tages in NPs synthesis. Fungal mycelium presents a specific external
surface and is resilient to shear forces and other unfavorable external
conditions (Fouda et al., 2017). Fungi excrete many extracellular
reductive enzymes which makes them good candidates for extracellular
NPs production (Chan and Mat Don, 2012; Narayanan and Sakthivel,
2010). Their ability to produce NPs is linked to their known metal ion
tolerance and bioaccumulation abilities (Dhillon et al., 2012; Sastry
et al., 2003). Another advantage of fungi mediated synthesis of NPs is
the creation of specific shapes and sizes and high degree of mono­
dispersion (Hulkoti and Taranath, 2014).
As with bacterial NPs synthesis, most published work concerns
AgNPs and AuNPs. However, other metal NPs synthesis by fungi and
yeasts has been reported (see Table 2). Cell-free filtrate of Aspergillus
fumigatus (Bhainsa and D’Souza, 2006) and Cryptococcus laurentii
(Fernández et al., 2016) has been used for AgNPs synthesis, as well as
the biomass of Fusarium oxysporum (Husseiny et al., 2015). AuNPs
synthesis by biomass cells has been reported for a wide variety of mi­
croorganisms, including F. oxysporum in a study by Mukherjee et al.
(2002) who achieved synthesis of spherical and triangular AuNPs by
extracellularly secreted reductases produced by precultured and washed
Fusarium oxysporum biomass which was combined with AuCl4− ions for
72 h in the dark. Other examples include Pichia jadinii (Gericke and
Pinches, 2006) and Trichoderma harzianum (Tripathi et al., 2018). Other
metal NPs synthesized by fungi include PdNPs (cell-free extract of
Saccharomyces cerevisiae (Sriramulu and Sumathi, 2018)) or PtNPs
(F. oxysporum biomass, (Riddin et al., 2006)).
2.2.3. Algae mediated NPs biosynthesis
The low media requirements, photosynthetic abilities and resistance
8
Biotechnology Advances 58 (2022) 107905
Applications Ref.
Antimicrobial (Abboud et al., 2014)
Catalytic (Rao and Gautam, 2016)
Catalytic (Khalafi et al., 2019)
Catalytic (Khalafi et al., 2019)
Antibacterial and antibiofilm (Ahiwale et al., 2017)
– (Love et al., 2015)
(Chen et al., 2015)
to high ion concentrations are the main advantages of algae, which can
be exploited in NPs synthesis. AgNPs synthesis by cell-free extracts was
reported by Moshfegh et al. (2019) for Polysiphonia sp.. Extracts from
Parachlorella kessleri has been studied for AgNPs synthesis (Kadukova
et al., 2014; Velgosova et al., 2020). Velgosova et al. (2020) studied the
biosynthesis of AgNPs resulting from three methods of algal extract
preparation. They reported that heating of the washed pre-cultured
algae, followed by centrifugation lead to optimal values of reaction
rate, uniformity and stability of NPs (~20 nm).
The biomass of Chlorella vulgaris (Luangpipat et al., 2011), Rhizo­
clonium fontinale (Parial and Pal, 2015) or Sargassum wightii (Singaravelu
et al., 2007) has been used for AuNPs synthesis. Other studies were
conducted with cell-free extracts of C. vulgaris for the synthesis of PdNPs
(Mishra et al., 2020) or ZnONPs (Khalafi et al., 2019).
2.2.4. Template mediated NPs biosynthesis
A novel approach to NPs synthesis is the use of biological templates
for the nanoparticle growth. Various molecules and structures can be
used as templates, e.g. fatty acids, amino acids, nucleic acids, capsules,
S-layers, and viroid or virus particles (Narayanan and Sakthivel, 2010).
The use of viruses as catalysts for NPs production is based on their
surface compounds protecting the capsid structure, which cooperate
specifically with metal ions (Salem and Fouda, 2021). With the use of
genetic engineering, self-assembled viral capsids were used for the
production of quantum dot NPs. In a study by Lee et al. (2002), a M13
bacteriophage with specific recognition moiety for ZnS crystal surfaces
was used to manufacture a self-supporting hybrid film material by
precipitation and suspension in ZnS precursor solution. Various metal
NPs, such as CdSNPs and PbSNPs were synthesized with the use of to­
bacco mosaic virus (Narayanan and Sakthivel, 2010; Shenton et al.,
1999).
3. Antimicrobial properties of NPs
One of the major challenges of modern medicine is the search for
new effective antimicrobial drugs, as many microbial pathogens are
developing resistance to classical antibiotics. In the medical field, the
potential of NPs has been studied in several applications, mostly as drug
carriers or parts of diagnostic systems, especially in cancer treatment.
However, their antimicrobial properties are also garnering attention
(Fahimmunisha et al., 2020; Luo et al., 2014; Salem and Fouda, 2021).
Various types of biosynthesized NPs were observed to possess antimi­
crobial activity against many pathogenic microorganisms and much
effort has thus been dedicated to evaluating their potential and ascertain
the mechanisms of action, which are summarized below in sections on
antibacterial (section 3.1), antifungal (section 3.2) and antiviral prop­
erties (section 3.3) and toxicity (section 4.5.3). For more details see
specialized reviews, e.g. (El-Seedi et al., 2019; Hassan et al., 2018;
Prabhu and Poulose, 2012; Salleh et al., 2020; Zhang et al., 2016a).
The antimicrobial activity of biosynthesized metal NPs has already
been observed against bacteria, fungi and even viruses, which are all
expanded on in following sections. Although there are many studies on
this topic, there is a great variability both in the NPs properties,
O. Mať átková et al.
microbial reactions and defenses and also in differing experimental
layout of various scientific studies.
There are several challenges that need to be addressed in the of green
synthesis studies regarding antimicrobial testing, both from the point of
experiment design and reporting and their future medicinal application.
Majority of the studies only use well or disc diffusion methods, which
does not constitute a comprehensive standard in microbiological
studies; the use of more advanced methods e.g., examining proteomics
or genomics, is scarce or non-existent. A lot of studies do not disclose
which strains of the microorganisms tested are used for the experiments
– this is crucial information in the context of microbiology field. Another
very prevalent shortcoming is non-sufficient method description, when
it comes to experiment layout, reagent composition or concentration of
nanoparticles used.
The physico-chemical properties of NPs like material, size (poly­
dispersity), shape, surface functionalization, etc. play a crucial role in
their antimicrobial effect, therefore it is important that the studies
include a thorough characterization of NPs and connect the microbio­
logical and physico-chemical properties. Smaller nanoparticles are
usually reported to have higher antimicrobial activity (Nisar et al., 2019;
Zhang et al., 2018). Their increased toxicity is hypothesized to results
from their higher specific surface area (compared to larger NPs) and
therefore a larger area available for the interaction with microbial cells
and for a quicker release of metal ions. However, there have been some
reports with opposing observations, i.e. that larger NPs had a better
antimicrobial effect. This suggests that the size is not the only factor that
affects nanoparticle activity (Slavin et al., 2017a). Another factor that is
known to affect the activity of a nanoparticle is shape. Cheon et al.
(2019) monitored the activity of silver nanoparticles of three different
shapes, namely, spherical, triangular plate, and disk, and found that
spherical nanoparticles had the highest activity. They found that the
different activity of the individual shapes is connected to the amount of
silver ions released from the surface (Cheon et al., 2019).
Moreover, it is often difficult to compare the antimicrobial properties
between individual studies due to great variability of methods, both in
the NPs synthesis and the microbiology evaluation. Antimicrobial ac­
tivity is usually evaluated by using well or disc agar diffusion methods,
growth curve analysis, changes in viability or total cell number (see
Table 3). Additionally, cell morphology and NPs localization or specific
metabolic activities are observed (Arora et al., 2008; Eid et al., 2020;
Salem and Fouda, 2021; Sayed and El-Sayed, 2020).
Several prospective mechanisms of NPs antimicrobial activity have
been suggested (see Fig. 4), depending on the microbial cell type and
particular metal NPs type. These are discussed below, however the main
mechanisms include damage to the cell membrane and wall, interactions
with intracellular components, and oxidative stress generation (Anjana
et al., 2019; Dikshit et al., 2021; Fang et al., 2018; Salem and Fouda,
2021).
In many cases, the NPs act through a combination of various mech­
anisms, which can be advantageous in medical applications, as the
probability of simultaneous resistance development is limited (Slavin
et al., 2017b). The advantage of biosynthesized NPs is in the function­
alization, capping, and stabilizing via biological molecules, which
themselves might possess antimicrobial properties and also allow the
NPs to bind to the target cell surface (Singh et al., 2018).
3.1. Antibacterial properties
Interactions of NPs with bacterial cells and their antibacterial
properties have been extensively studied in a variety of metal NPs and
strains of both Gram-positive and Gram-negative bacteria (see Table 3).
Beside the innate antimicrobial activity of biosynthesized NPs, they
have been reported to facilitate enhancement of the activity of con­
ventional antibiotics, e.g. AgNPs were proven effective in combination
with levofloxacin against Enterococcus faecalis, Staphylococcus aureus,
E. coli, Proteus vulgaris, Salmonella typhi, and Vibrio cholerae (Keshari
9
Biotechnology Advances 58 (2022) 107905
et al., 2020). The most reported biosynthesized NPs with antibacterial
properties are silver, gold, palladium, copper and copper oxide, zinc,
and zinc oxide (Das et al., 2017; Gericke and Pinches, 2006; Narayanan
and Sakthivel, 2010).
3.1.1. Silver NPs
The application of antibacterial properties of AgNPs has been re­
ported in several reviews, both prepared by physico-chemical and bio­
logical methods (Prabhu and Poulose, 2012; Rafique et al., 2017; Tran
et al., 2013; Zhang et al., 2016b). The main mechanisms of action of
AgNPs are the release of Ag+ ions, ROS generation, interference with
energy metabolism and cell wall synthesis, membrane disruption and
interference with DNA, RNA and other parts of cell metabolism (Dakal
et al., 2016; Gurunathan et al., 2014; Lara et al., 2010b; Singh et al.,
2018). These are facilitated both by the extracellular and intracellular
presence of AgNPs, as small sized AgNPs can penetrate the cell wall and
membrane. Their transport through cell wall may be hindered by the
peptidoglycan layer in Gram-positive bacteria, resulting in their lower
susceptibility in some studies (Gurunathan et al., 2014). Another cor­
relation has been found with the size and shape of NPs, as smaller NPs
(higher surface to volume ratio) have shown quicker release of Ag+ ions
(Tang and Zheng, 2018). Similarly, it was found that spherical AgNPs
exhibit higher antimicrobial activity than their triangular, platelet or
nanorod counterparts because of enhanced surface binding and uptake
(Helmlinger et al., 2016).
Bacterial species which have been reported as susceptible to bio­
synthesized AgNPs include multidrug resistant pathogens, such as Ba­
cillus subtilis, E. coli, Enterococcus sp., Klebsiella pneumoniae, Proteus
mirabilis, P. aeruginosa, S. aureus, Shigella flexaneri and V. cholerae
(Ahmed et al., 2016; Nayak et al., 2016; Składanowski et al., 2017).
Synergistic effects of AgNPs with conventional antibiotics (vancomycin,
clindamycin, amoxicillin, erythromycin) have been shown against
S. aureus and E. coli (Shahverdi et al., 2007; Singh et al., 2018).
3.1.2. Gold NPs
Biosynthesized AuNPs have also been widely studied, although their
antibacterial activities are limited in comparison with AgNPs. AuNPs are
a target of studies of biocomposites, where the AuNP serves as a carrier
for an antibiotic, which showed activity against multi-drug resistant
bacteria including methicillin resistant S. aureus, overcoming the beta-
lactamase based resistance of these strains (Brown et al., 2012). How­
ever, AuNPs alone have been reported to act at least bacteriostatically
against a variety of bacterial strains (E. coli, B. subtilis, K. pneumoniae)
(Singh et al., 2018). There are two main proposed mechanisms of action
of AuNPs: inhibiting ATPase activity, which hinders energy metabolism,
and preventing tRNA ribosome interaction, which stops transcription
(Cui et al., 2012).
3.1.3. Metal oxide NPs
Antibacterial activity has been found in biosynthesized ZnONPs
against a wide variety of microorganisms including carbapenem resis­
tant Acinetobacter baumanii (Tiwari et al., 2018). This activity is con­
nected to ROS generation, cell membrane damage, and interactions of
both ZnONPs and Zn ions with intracellular components (Saravanan
et al., 2018b; Singh et al., 2018). Although biological synthesis of
CuONPs is a new topic, interesting mainly due to their resistance to
oxidation reactions which are likely to occur in biological media, there
have been reports on their antibacterial activity against both Gram-
positive and Gram-negative bacteria (DeAlba-Montero et al., 2017).
MgONPs activity is reportedly based on the generation of ROS and
direct interactions with the cell membrane (Nguyen et al., 2018);
however, some reports have demonstrated antibacterial activity without
ROS-induction (Leung et al., 2014). Antibacterial effect of TiO2NPs that
has been demonstrated against various microorganisms is also con­
nected with ROS generation, in this case induced by their photosensi­
tivity in the near UV spectrum (Li et al., 2012b; Singh et al., 2018).
O. Mať átková et al. Biotechnology Advances 58 (2022) 107905

Table 3
Antimicrobial properties of biosynthesized metal NPs against various microorganisms.
Target microorganism NP type, size (nm) Biological agent used for Method Ref.
synthesis

Bacteria
Acinetobacter sp. Ag, 5–20 Chlorella pyrenoidosa Well diffusion (Aziz et al., 2015)
Aeromonas hydrophila Ag, 5–20 Chlorella pyrenoidosa Well diffusion (Aziz et al., 2015)
Bacillus cereus Ag, - Ananas comosus Well diffusion (Das et al., 2019)
Enterococcus faecium Ag, - Ananas comosus Well diffusion (Das et al., 2019)
Enterococcus faecalis Ag, 5–50 Rhodococcus sp. Growth curve, well diffusion (Otari et al., 2015)
Enterobacter aerogenes ZnO, 15–30 Streptomyces sp. Well diffusion (Shanmugasundaram and
Balagurunathan, 2017)
Escherichia coli Ag, <24 Impatiens balsamina, Lantana Disc diffusion (Aritonang et al., 2019)
camara
Disc diffusion, Congo red agar, crystal violet
Ag, 20 Solanum nigrum (Jinu et al., 2017)
staining
Ag, 5–50 Rhodococcus sp. Growth curve, well diffusion (Otari et al., 2015)
Ag, 7–28 Streptomyces laurentii Well diffusion (Eid et al., 2020)
Ag, 16–70 Fusarium oxysporum Turbidimetric assay (Gopinath et al., 2015)
OD measurement, high-content screening,
Ag, 3–15 Chlorella vulgaris (da Silva Ferreira et al., 2017)
ultrastructural analysis
CuO, 2–41 Bacillus sp. Disc diffusion (Taran et al., 2017)
CuO, 20–40 Solanum lycopersicum Well diffusion, SEM (Vaidehi et al., 2018)
Pd, 15–90 Rosmarinus officinalis Well diffusion (Rabiee et al., 2020)
(Shanmugasundaram and
ZnO, 15–30 Streptomyces sp. Well diffusion
Balagurunathan, 2017)
ZnO, 9–22 Fusarium solani Disc diffusion (Sayed and El-Sayed, 2020)
Disc diffusion, Congo red agar, crystal violet
Ag, 20 Solanum nigrum (Jinu et al., 2017)
staining
Ag, 5–50 Rhodococcus sp. Growth curve, well diffusion (Otari et al., 2015)
Ag, 16–70 Fusarium oxysporum Turbidimetric assay, well diffusion (Gopinath et al., 2015)
OD measurement, high-content screening,
Klebsiella pneumoniae Ag, 3–15 Chlorella vulgaris (da Silva Ferreira et al., 2017)
ultrastructural analysis
Ag, 2–15 Candida glabrata MIC, MBC (Jalal et al., 2018)
(Shanmugasundaram and
ZnO, 15–30 Streptomyces sp. Well diffusion
Balagurunathan, 2017)
ZnO, 9–22 Fusarium solani Disc diffusion (Sayed and El-Sayed, 2020)
Listeria monocytogenes Ag, - Ananas comosus Well diffusion (Das et al., 2019)
Micrococcus lutens Pd, 15–90 Rosmarinus officinalis Well diffusion (Rabiee et al., 2020)
Disc diffusion, Congo red agar, crystal violet
Ag, 20 Solanum nigrum (Jinu et al., 2017)
staining
Proteus vulgaris Ag, 5–50 Rhodococcus sp. Growth curve, well diffusion (Otari et al., 2015)
(Shanmugasundaram and
ZnO, 15–30 Streptomyces sp. Well diffusion
Balagurunathan, 2017)
Ag, - Lantana camara Well diffusion (Patil Shriniwas, 2017)
Disc diffusion, Congo red agar, crystal violet
Ag, 20 Solanum nigrum (Jinu et al., 2017)
staining
Ag, 5–50 Rhodococcus sp. Growth curve, well diffusion (Otari et al., 2015)
Pseudomonas
Ag, 7–28 Streptomyces laurentii Well diffusion (Eid et al., 2020)
aeruginosa
Ag, 2–15 Candida glabrata MIC, MBC (Jalal et al., 2018)
Ag, 16–70 Fusarium oxysporum Turbidimetric assay, well diffusion (Gopinath et al., 2015)
CuO, 40–110 Lactobacillus casei MIC, MBC, well diffusion (Kouhkan et al., 2020)
ZnO, 9–22 Fusarium solani Disc diffusion (Sayed and El-Sayed, 2020)
Pseudomonas
Ag, 10–20 Bacillus subtilis MIC, MBC (Velmurugan et al., 2014)
fluorescens
Ag, 2–15 Candida glabrata MIC, MBC (Jalal et al., 2018)
Salmonella enterica (Shanmugasundaram and
ZnO, 15–30 Streptomyces sp. Well diffusion
Balagurunathan, 2017)
Shigella flexneri Ag, 2–15 Candida glabrata MIC, MBC (Jalal et al., 2018)
Ag, - Ananas comosus Well diffusion (Das et al., 2019)
Impatiens balsamina, Lantana
Ag, <24 Disc diffusion (Aritonang et al., 2019)
camara
Ag, - Lantana camara Well diffusion (Patil Shriniwas, 2017)
Disc diffusion, Congo red agar, crystal violet
Ag, 20 Solanum nigrum (Jinu et al., 2017)
staining
Ag, 10–20 Bacillus subtilis MIC, MBC (Velmurugan et al., 2014)
Ag, 13–28 Lactobacillus plantarum Well diffusion (Garmasheva et al., 2016)
Ag, 5–50 Rhodococcus sp. Growth curve, well diffusion (Otari et al., 2015)
Staphylococcus aureus
OD measurement, high-content screening,
Ag, 3–15 Chlorella vulgaris (da Silva Ferreira et al., 2017)
ultrastructural analysis
Ag, 4–50 Sinomonas mesophila Well diffusion (Manikprabhu et al., 2016)
Ag, 2–15 Candida glabrata MIC, MFC (Jalal et al., 2018)
CuO Bacillus sp. Disc diffusion (Taran et al., 2017)
CuO, 20–40 Solanum lycopersicum Well diffusion, FE-SEM (Vaidehi et al., 2018)
CuO, 40–110 Lactobacillus casei MIC, MBC, well diffusion (Kouhkan et al., 2020)
Pd, 15–90 Rosmarinus officinalis Well diffusion (Rabiee et al., 2020)
ZnO, 15–30 Streptomyces sp. Well diffusion
(continued on next page)

10
O. Mať átková et al. Biotechnology Advances 58 (2022) 107905

Table 3 (continued )
Target microorganism NP type, size (nm) Biological agent used for Method Ref.
synthesis

(Shanmugasundaram and
Balagurunathan, 2017)
ZnO, 9–22 Fusarium solani Disc diffusion (Sayed and El-Sayed, 2020)
Disc diffusion, Congo red agar, crystal violet
Ag, 20 Solanum nigrum (Jinu et al., 2017)
Staphylococcus staining
epidermidis Pd, 15–90 Rosmarinus officinalis Well diffusion (Rabiee et al., 2020)
Ag, 13–28 Lactobacillus plantarum Well diffusion (Garmasheva et al., 2016)
Fungi and yeasts
Alternaria alternata CuO, - Streptomyces zaomyceticus Well diffusion (Hassan et al., 2019)
Cu / ZnO, 10–20 /
Aspergillus awamori Fusarium solani Disc diffusion (Sayed and El-Sayed, 2020)
9–22
Cu / ZnO, 10–20 /
Aspergillus fumigatus Fusarium solani Disc diffusion (Sayed and El-Sayed, 2020)
9–22
Aspergillus niger CuO, - Streptomyces zaomyceticus Well diffusion (Hassan et al., 2019)
Ag, 13–28 Sinomonas mesophila Well diffusion (Manikprabhu et al., 2016)
Candida albicans Ag, 2–15 Candida glabrata MIC, MFC, ultrastructural analysis (Jalal et al., 2018)
Pd, 15–90 Rosmarinus officinalis Disc diffusion (Rabiee et al., 2020)
Ag, 2–15 Candida glabrata MIC, MFC, ultrastructural analysis (Jalal et al., 2018)
Candida glabrata
Pd, 15–90 Rosmarinus officinalis Disc diffusion (Rabiee et al., 2020)
Ag, 2–15 Candida glabrata MIC, MFC, ultrastructural analysis (Jalal et al., 2018)
Candida krusei
Pd, 15–90 Rosmarinus officinalis Disc diffusion (Rabiee et al., 2020)
Ag, 2–15 Candida glabrata MIC, MFC, ultrastructural analysis (Jalal et al., 2018)
Candida parapsilosis Ag, 5–45 Rhodotorula glutinis Disc diffusion (Cunha et al., 2018)
Pd, 15–90 Rosmarinus officinalis Disc diffusion (Rabiee et al., 2020)
Cu / ZnO, 10–20 /
Fusarium solani Disc diffusion (Sayed and El-Sayed, 2020)
Fusarium oxysporum 9–22
CuO, - Streptomyces zaomyceticus Well diffusion (Hassan et al., 2019)
Pythium ultimum CuO, - Streptomyces zaomyceticus Well diffusion (Hassan et al., 2019)

(MIC – minimum inhibitory concentration, MFC – minimum fungicidal concentration, MBC – minimum bactericidal concentration, OD – optical density)

Fig. 4. Antimicrobial mechanisms of metal NPs application affecting intracellular components and cell interface.

3.1.4. Mechanisms of activity of metal NPs
The interactions of metal NPs are both varied and individual to
specific metal and microorganism; however, as described above, most
metal NPs act in several different ways, which is one of the main attri­
butes of their potential medical application. The most frequently pro­
posed antibacterial activity mechanisms of metal NPs are oxidative
stress related to ROS production, metal ion release, and various non-
oxidative mechanisms. The non-oxidative mechanisms which were re­
ported include direct interactions with cell wall and membrane, pene­
tration into cell interior and subsequent inhibition of nucleic acid and
protein synthesis and regulation of gene expression. Metal NPs are also
known to interact with and disrupt bacterial biofilm (Gurunathan et al.,
2014), thus hindering an important resistance mechanism which pre­
sents the bacterial cells with a significant advantage.
Most of these mechanisms are executed through direct contact of the
NPs with the bacterial cell, which is enabled by electrostatic in­
teractions, van der Waals forces, and hydrophobic interactions (El-Seedi
et al., 2019; Slavin et al., 2017b). Penetration through the cell wall and
membrane is size-dependent, as are the subsequent interactions with
intracellular cell components, such as proteins, nucleic acids, ribosomes,
lysosomes, etc. (Mukha et al., 2013). These interactions can cause
oxidative stress, enzyme deactivation, modification of gene expression,
alteration of cell membrane permeability and electrolyte imbalance
(Wang et al., 2017). The mechanisms of NPs penetration into the cell
interior include depolarization of cell wall, which subsequently becomes
more permeable, ability to cause irregular pit formations on the cell
wall, consequently leading to cell wall degradation, receptor-mediated
endocytosis and diffusion through the pores in the cell membrane (Lin

11
O. Mať átková et al.
et al., 2010; Prabhu and Poulose, 2012; Slavin et al., 2017b).
Oxidative stress induced by ROS production is one of the most
studied antibacterial mechanisms of NPs. Different metal NPs produce
various ROS compounds, producing a wide range of specific antibacte­
rial activities (Wang et al., 2017). Excessive ROS production leads to
oxidative stress for the bacterial cell, which is followed by loss of cell
membrane integrity, interaction with DNA and enzymes and triggering
cell apoptosis (Cheloni et al., 2016; Pramanik et al., 2012).
The physico-chemical properties of NPs are crucial in the determi­
nation of the possible mechanisms of their antibacterial action, among
the most important are their size, shape, roughness, specific surface
area, surface energy, charge, zeta potential, surface morphology, crystal
structure and ligands (Wang et al., 2017). The activity of metal NPs is
also influenced by the conditions in which the antibacterial activity is
studied. The medium composition, pH, osmotic pressure, temperature,
and irradiation can influence properties of NPs like solubility, aggre­
gation and surface charge or modification, which can all influence the
observed antimicrobial properties reported in literature (Wang et al.,
2017).
3.2. Antifungal properties
Metal NPs have been used as potential inhibitors of fungal patho­
genic microorganisms, with more focus on NPs obtained by physical and
chemical methods than on their biosynthesized counterparts (Sousa
et al., 2020; Tran et al., 2013). Their mechanisms of activity are stated as
consistent with antibacterial properties, i.e. oxidative stress via ROS
production, cell wall and membrane damage due to direct contact with
NPs, cellular uptake of NPs and subsequent damaging interactions with
cell interior and metal ion release (Aljabali et al., 2018; Sousa et al.,
2020).
Gupta et al. (2014) reported antifungal activity of biosynthesized
AgNPs against Candida albicans, Trichophyton mentagrophytes, and
Microsporum canis, both alone and in synergism with antibiotics.
AgClNPs proved antifungal activity against C. albicans, Aspergillus flavus,
and Aspergillus niger (Paulkumar et al., 2013).
Moreover, studies have also shown the antifungal activity of green
AuNPs synthesized via both plant and microbial synthesis. This inhibi­
tion of fungi by biosynthesized AuNPs has already been reported against
Puccinia graminis tritci, Microphyton gypseum, A. flavus, A. niger, Tricho­
phyton rubrum and C. albicans (Gopal et al., 2013).
3.3. Antiviral properties
The search for new and effective antiviral agents is an essential part
of modern medicine. Although the antiviral abilities of NPs synthesized
by physical and chemical methods are known (Lara et al., 2010a; Xiang
et al., 2011), even here the exact mechanism of their antiviral activity is
an ongoing research topic, with proposed mechanisms including inhi­
bition of virion formation, preferential binding preventing host cell
attachment and inhibition of viral particles replication (Zhang et al.,
2016b). The antiviral properties of biosynthesized AgNPs were reported
against SARS-CoV-2 (Al-Sanea et al., 2021), HIV type 1 (Kumar et al.,
2017), herpes simplex virus (type 1 and 2), and human parainfluenza
virus type 3 (Gaikwad et al., 2013). Ag2O/AgONPs and AuNPs showed
reduction activity of the cytopathic effect of the herpes simplex virus
(type 1) (El-Sheekh et al., 2020).
4. Application and limitations of NPs
Due to their unique properties and advantages, metal NPs are finding
applications in various medicinal, biotechnological and industrial areas.
In addition, the biosynthesized NPs offer their biocompatibility and
economical and ecological benefits of green synthesis and thus further
broaden their possible applications. NPs are applicable in many tradi­
tional and novel emerging technologies, both as substitutes and as part
12
Biotechnology Advances 58 (2022) 107905
of novel approaches (Ahmed et al., 2016; El-Seedi et al., 2019; Salem
and Fouda, 2021), their application potential in various areas is
described below, namely, medicine (section 4.1), agriculture (4.2), in­
dustry (4.3), water-treatment and bioremediation (4.4).
4.1. Application of NPs in medicine
The development of novel drugs, treatments and diagnostic systems
is an integral part of modern medicinal research. The ongoing research
on green synthesized NPs for medical applications focuses on NPs from
various material, especially metals and their oxides, such as Ag, Au,
ZnO, TiO2, CuO, MgO, and SiO2 (Dikshit et al., 2021; Ge et al., 2014;
Nath and Banerjee, 2013). Their main potential lies in imaging systems,
biosensors, drug-delivery and treatment for multi drug resistant mi­
crobes (Fahimmunisha et al., 2020; Salem and Fouda, 2021).
Further potential can be found in the synthesis of nanomaterials and
nanocomposites utilizing beneficial properties of NPs with other me­
dicinal materials, e.g. for immunoassays and antimicrobial combination
therapy (Fayaz et al., 2010; Khani et al., 2018; Le et al., 2017). The use of
NPs in cancer treatment and diagnostics utilizes their large surface area
and uses them as carriers in combination with targeted therapy. This
approach poses additional benefits such as little negative impact,
effectivity and biocompatibility (Salem and Fouda, 2021).
Medical materials like wound dressings with traditionally synthe­
sized AgNPs have been developed and used in clinical environment
(Chaloupka et al., 2010) and were proven to reduce healing time in some
wound types, possibly due to their antimicrobial properties (Huang
et al., 2007). The biosynthesized AgNPs might offer further advantages
regarding their higher biocompatibility. Similarly, there are silver-
coated implants (catheters, cardiovascular implants), which might
benefit from integrating AgNPs, either as coatings or in nanocomposite
materials (Ge et al., 2014).
4.2. Application of NPs in agriculture
Modern agriculture benefits from the use of new technologies,
including the use of NPs. Their potential lies mainly in pest control and
targeted nutrient delivery (Salem and Fouda, 2021).
The NPs can be part of nanopesticide formulations, targeted at mi­
crobial pathogens or insects. Wang et al. (2007) proposed emulsion-
based products containing NPs that proved to be effective against a
variety of agricultural pests.
The potential of biosynthesized NPs as antifungal agents for common
plant pathogens like Aspergillus sp., Fusarium sp. and Pythium sp. has
been demonstrated by Hassan et al. (2018). Biosynthesized SeNPs were
proposed as possible control agents for malaria vector Anopheles ste­
phensi (Salem et al., 2021).
Beside the control of pests, the efficient use of resources and coun­
teracting water and soil fertility decline is another goal of sustainable
agriculture and a possible area for NPs application. These include stra­
tegies for slow release of various fertilizers, e.g. by implementing
nanocoating (Duhan et al., 2017).
4.3. Application of NPs in industry
The application of NPs in textile industry has increased in recent
years, as they were found to provide beneficial properties to the pro­
duction process and the fabrics (Salem and Fouda, 2021). Beside the
antimicrobial properties of various NPs (Ag, ZnO), they also add UV
blocking and self-cleaning attributes to the final product (Fouda et al.,
2018; Fouda et al., 2017). They provide excellent chemical stability and
the large surface to volume ratio provides significant advantages over
organic counterparts when it comes to UV blocking efficiency (Shaheen
et al., 2019).
In food industry, the use of nanomaterials as additivities or coating of
materials can provide significant benefits in food safety and packaging
O. Mať átková et al.
due to their antimicrobial and antibiofilm properties. The use of pack­
aging or containers with incorporated NPs aims at food contamination
prevention and increased resistance to microbial colonization (Sharma
et al., 2017).
The optical properties of NPs, nanocomposite materials especially,
can be exploited in functional thin-film coatings, e.g. by thermal treat­
ment of nanocrystalline AgNPs and formation of carbonaceous nano­
materials (Klaus et al., 1999).
The biosynthesis of nanoscale semiconductors has been studied with
potential application both in electrotechnical industry and for their
optical properties (Mandal et al., 2006). Nanoparticles composed of CdS,
ZnS, and PbS are studied for their luminescent properties in biological
imaging as quantum dots. The conjugation of semiconductor NPs with
specific biomolecules via biosynthetic route can result in a wide variety
of quantum dots with different chemical compositions and sizes, and
therefore broaden their potential applications, such as cell labelling,
imaging, compound tracking or in biosensors (Valizadeh et al., 2012).
The biosynthesized semiconductor NPs such as CdS crystallites have also
been studied for their quantum properties aiming at overcoming limits
in microelectronics (Dameron et al., 1989).
Copper and copper oxide NPs have many applications in the con­
struction of modern electronic circuits due to their conductive, elec­
tronic and optical properties and have also been reported in catalysis
applications (Gawande et al., 2016).
A range of nanomaterials with novel properties can be obtained by
using NPs as input material for further modification and treatment.
Using isolated NPs or even combining biosynthetic methods com­
plemented by classical preparation methods and materials such as ce­
ments, ceramics and plastics can yield novel hybrid and composite
materials, such as metal-organic frameworks or graphene-metal com­
posites (Kharissova et al., 2019).
Magnetic NPs can aid in synthesis of industrial catalysts which are
easily recoverable and recyclable. Their use has been developed for
various reductive, oxidative or condensation reactions and pose ad­
vantages in high reaction times due to the nanocatalytic properties and
ease of the separation due to the magnetic properties of the material
after each reaction cycle (Varma, 2012).
4.4. Application of NPs in water treatment and bioremediation
Sustainable water management and treatment is considered one of
the most urgent problems of modern society. Nanotechnology including
nanomaterials can provide efficient systems for the removal of patho­
gens, pollutants, toxic compounds and heavy metals (Uddandarao et al.,
2019; Zonaro et al., 2017). The proposed systems include functionalized
nanofiber membranes or membranes with associated immobilized NPs.
In wastewater treatment the NP-associated membrane aids in pollutant
removal based on combination of adsorption, filtration and degradation
mechanisms. Such membrane modules can be included in both classical
wastewater treatment plants or in ex situ bioreactor bioremediation
techniques (El-Aswar et al., 2022). Bimetallic NPs, prepared using green
tea extracts and associated with membranes have been used in the
degradation of trichloroethylene (Varma, 2012).The feasibility of these
systems has been demonstrated for ZnONPs in pathogenic microbe or
metal removal from wastewater (Primo et al., 2020; Rambabu et al.,
2021) or CuONPs in toxic dye removal from wastewater (Rafique et al.,
2020). The catalytic properties of metal NPs can be used in customized
waste treatment and bioremediation processes, as both AgNPs and
AuNPs have been shown to catalyze degradation of organic dyes and
other pollutants (Salem and Fouda, 2021; Suvith and Philip, 2014).
Bioremediation applications using reductive properties of nanoscale
zero-valent iron (NZVI) towards pollutants have been applied as
permeable reactive barriers or direct injection in situ (Mueller et al.,
2012). Plant waste (such as sorghum bran) derived iron NPs combined
with natural surfactants have been reported as posing a stable, envi­
ronmentally friendly, easily stored material usable for catalytic
13
Biotechnology Advances 58 (2022) 107905
activation of hydrogen peroxide and remediation of contaminated soils
(Varma, 2012).
4.5. Limitations and safety of biosynthesized NPs
Beside the many benefits and advantages of NPs, there are many
considerations that need to be discussed with respect to the possible
widespread use in future. The limitations arise both in the process of
manufacture and in the guarantee of safety for humans and their fate in
the environment.
4.5.1. Green synthesis
The green methods of synthesis, although by principle more con­
scientious toward waste management and other environmental con­
cerns, are not yet as advanced and widespread. The large-scale
manufacture of NPs by green methods needs case by case optimization,
with respect to the desired properties of NPs (shape, size, biological
activity). Among the reported challenges are detailed knowledge of the
synthesis mechanisms with the roles of various compounds, uniformity
of the biological material and studies focused on the procedures, econ­
omy and waste management of the scaled-up processes (Salem and
Fouda, 2021).
The assessment of the sustainability of green methods in synthesis of
metal NPs is a key element in their long-term applicability and evalua­
tion. This can be aided by the use of modelling including set of decision
rules which can encompass key variables and target evaluations,
including life cycle analysis, synthesis protocols, including identification
of key variable conditions and allowing for overall assessment and
comparison between techniques and approaches. Cinelli et al. (2015)
developed a modelling approach for this purpose, allowing selecting
criteria, evaluation and classification of AgNPs synthesis methods and
protocols (based on green chemistry) into performance classes, as well
as their robustness (Cinelli et al., 2017; Cinelli et al., 2015). The eval­
uation of the sustainability of the proposed NPs synthesis protocols or
developments of industrial-scale methods can be included in the models
as a set of environmentally oriented criteria which aim at aiding decision
planning while incorporating both experimental results and reference
protocols. Multiple criteria analysis in a co-constructive development of
a classification model for NPs synthesis assessment model has been
proposed by Kadziński et al. (2018). Life cycle analysis has revealed that
the choice of synthesis route for AgNPs production has a great impact on
the energy use and environmental impacts; however, the goals (with
respect to size and antimicrobial efficacy) significantly influence the
performance-based comparison (Pourzahedi and Eckelman, 2015).
4.5.2. NPs in environment
Although the biosynthesized NPs should present lesser toxicity then
those synthesized by traditional methods, the impacts of NPs release into
the environment and the effects on ecosystem balance need to be thor­
oughly studied (de Souza et al., 2019). The fate of NPs in the environ­
ment includes the direct effect of the original NPs and their
transformation products, which all need to be considered (Levard et al.,
2012).
An important aspect of the evaluation of NPs impact of environment
is their origin. The toxicity of naturally occurring nanoparticles can
differ significantly from those of synthetic origin. Naturally formed NPs
(metals and their oxides and sulfides) can occur in aquatic, atmospheric,
lithospheric and biospheric environments, e.g. in wastewaters, around
hydrothermal vents or in ore deposits. They can be formed via physio-
chemical, mechanical, and biological routes under variety of environ­
mental conditions including thermal, non-thermal, and photochemical
conditions which are influenced by various environmental factors (pH,
temperature, oxygen, presence of ROS, ions and organic compounds of
natural origin etc.). Their fate and toxicity is also significantly influ­
enced by the external conditions during their formation and afterwards,
as the functional groups derived from natural organic matter capping
O. Mať átková et al.
agents strongly affect their possible interactions with the environment
and its organisms (Sharma et al., 2015).
The direct influence of metal NPs depends on the type; however, it
can include the interactions with biological matter, living organisms,
and inorganic components of the environment. The antimicrobial
properties of NPs can negatively affect autochthonous microflora of the
given ecological niche, including the spread of resistance (Miller et al.,
2010). Unutilized waste NPs can become widespread in nature and there
are some studies concerning their presence in environment, including
food and water, with unknown impacts (Tiede et al., 2008).
The transformation reactions of NPs in the environment involve
dissolution and release of ions (especially in AgNPs), oxidative or
reductive reactions, aggregation, and adsorption (de Souza et al., 2019).
These processes are dependent on the NPs type, size and shape, their
capping agents, but also on the environmental conditions, like pH,
temperature, ionic strength and presence of reactive compounds
(Sharma and Zboril, 2017; Zhang et al., 2016b). The aggregation and
adsorption processes are highly dependent on the type of capping agents
(biological compounds in biosynthesized NPs) and their possible in­
teractions with the environment, which can both facilitate and hinder
the transformation processes (de Souza et al., 2019; Grillo et al., 2015).
As these processes are both highly individual and also all act
simultaneously and interfere with each other, more research is needed to
evaluate the impact of potential widespread use of NPs.
4.5.3. NPs toxicity and biological compatibility
The toxicity of NPs in the environment is highly dependent on the NP
type, size and shape and on the transformations it undergoes. In AgNPs,
the oxidative dissolution is responsible for release of Ag+ ions which are
highly toxic to many organisms even in low concentrations which can
possibly occur in the environment (Ašmonaitė et al., 2016; Jiravova
et al., 2016). The exposure and residence time of NPs in the environment
can be significantly prolonged by their adsorption onto organic or
inorganic matter, thus increasing their persistence (Grillo et al., 2015).
In some cases, the adsorption can mitigate the toxicity of NPs by
complex-formation of stabilizing reactions, which alleviate the effect on
the environment (Kim et al., 2013). The biosynthesized NPs have been
observed to possess lesser toxicity, which is generally contributed to the
use of biological components as capping agents and stabilizers, as well as
avoidance of toxic solvents during their manufacture (Hasan et al.,
2020; Mražiková et al., 2017). Non-toxicity of biosynthesized iron oxide
nanoparticles (at 2 g/kg) was demonstrated on laboratory mice in a
study conducted by Lakshmi Pravallika et al. (2019).
The application of NPs in medicine is dependent on the minimization
of potential NPs side-effects and their impact on the organism. The in­
teractions between NPs and host tissues are dependent on their prop­
erties, such as material, size and shape (Li et al., 2012a; Rudramurthy
et al., 2016). The majority of NPs have the ability to permeate cell
membranes and travel through the blood stream and thus spread to
various parts of the body. They have even been demonstrated to be able
to cross the blood-brain barrier (Oberdorster et al., 2009; Rudramurthy
et al., 2016). Potential harm also lies in their selective accumulation in
specific tissues (Buzea et al., 2007).
Interactions with cells and intracellular components can have
adverse effects and cause cellular damage, e.g. via oxidative stress. The
size of NPs seems to be the determining factor in the mitigation of their
spread through the organism and the subsequent immune system. The
oxidative stress resulting from NPs presence can contribute to inflam­
mation, toxicity, DNA and mitochondrial damage and thus to the
development of serious conditions like pulmonary inflammation or
kidney and liver issues (Dikshit et al., 2021).
The biological effects and compatibility with living organisms and
tissues for specific medicinal application need to be established before
their application. Evaluation criteria can be established with respect to
in vitro cytotoxicity tests, hemolysis, blood cell aggregation, neurotox­
icity, immunotoxicity, histological compatibility and other attributes
14
Biotechnology Advances 58 (2022) 107905
(Pourali et al., 2017; Rudramurthy et al., 2016).
More research is needed to evaluate the potential adverse effects of
any medicinal NPs application and to establish all potential limitations
in an application-dependent manner.
5. Concluding remarks
Increasing industrial demands and environmental concerns have
given rise to the development of green methods for the synthesis of
metal nanoparticles. The potential of biologically synthesized nano­
particles potential lies in the eco-friendly and cost-effective methods and
materials utilized for their synthesis and in their biological compatibility
due to the use of biological molecules as stabilizers and capping agents.
Plant and microbial organisms and extracts offer a wide variety of agents
suitable for green methods. This variety offers many benefits in the
flexibility and uniqueness of the final nanoparticles properties but also
presents a challenge for the scientific community to ascertain all the
underlying molecular mechanisms in the synthesis and to evaluate all
the characteristics and potential limitations of the final nanoparticles.
The potential of nanoparticles is especially valuable in medical appli­
cations, due to the high effectiveness of metal nanoparticles against a
variety of serious human pathogens, including multidrug resistant mi­
croorganisms. Other potential areas of application include agriculture
and food industry. This broad potential of nanoparticles application
brings an urgent need for a deep understanding of all aspects of nano­
particles interaction with human body and with the environment and
their overall impact on a local and global scale.
Author contributions
The author contribution statement is based on the CReditT role
system.
Funding
This work was supported by the Technology Agency of the Czech
Republic [project TH04030109].
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