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For purpose of this review, silver nanomaterials include silver nanoparticles, stabilized silver salts, silver–dendrimer, polymer and metal oxide composites, and silver-impregnated zeoli

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R E V I E W P A P E R

A review of the antibacterial effects of silver nanomaterials

and potential implications for human health

and the environment

Catalina Marambio-Jones•Eric M V Hoek

Received: 29 July 2009 / Accepted: 6 March 2010 / Published online: 23 March 2010

Ó Springer Science+Business Media B.V 2010

Abstract Here, we present a review of the

antibac-terial effects of silver nanomaantibac-terials, including

pro-posed antibacterial mechanisms and possible toxicity

to higher organisms For purpose of this review, silver

nanomaterials include silver nanoparticles, stabilized

silver salts, silver–dendrimer, polymer and metal oxide

composites, and silver-impregnated zeolite and

acti-vated carbon materials While there is some evidence

that silver nanoparticles can directly damage bacteria

cell membranes, silver nanomaterials appear to exert

bacteriocidal activity predominantly through release of

silver ions followed (individually or in combination)

by increased membrane permeability, loss of the

proton motive force, inducing de-energization of the

cells and efflux of phosphate, leakage of cellular

content, and disruption DNA replication Eukaryotic

cells could be similarly impacted by most of these

mechanisms and, indeed, a small but growing body of

literature supports this concern Most antimicrobial

studies are performed in simple aquatic media or cell

culture media without proper characterization of silver

nanomaterial stability (aggregation, dissolution, and

re-precipitation) Silver nanoparticle stability is

gov-erned by particle size, shape, and capping agents as

well as solution pH, ionic strength, specific ions andligands, and organic macromolecules—all of whichinfluence silver nanoparticle stability and bioavailabil-ity Although none of the studies reviewed definitivelyproved any immediate impacts to human health or theenvironment by a silver nanomaterial containingproduct, the entirety of the science reviewed suggestssome caution and further research are warranted giventhe already widespread and rapidly growing use ofsilver nanomaterials

Keywords Silver Nanoparticle Antimicrobial  Antibacterial  Nanotechnology Nanotoxicology Safety  EHS

IntroductionThe broad-spectrum antimicrobial properties of silverencourage its use in biomedical applications, waterand air purification, food production, cosmetics,clothing, and numerous household products Withthe rapid development of nanotechnology, applica-tions have been extended further and now silver is theengineered nanomaterial most commonly used inconsumer products (Rejeski 2009) Clothing, respi-rators, household water filters, contraconceptives,antibacterial sprays, cosmetics, detergent, dietarysupplements, cutting boards, sox, shoes, cell phones,laptop keyboards, and children’s toys are among the

C Marambio-Jones  E M V Hoek (&)

Department of Civil and Environmental Engineering,

California NanoSystems Institute, University

of California, Los Angeles, 5732G Boelter Hall,

PO Box 951593, Los Angeles, CA 90095-1593, USA

e-mail: emvhoek@ucla.edu

DOI 10.1007/s11051-010-9900-y

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retail products that purportedly exploit the

antimi-crobial properties of silver nanomaterials

Different forms of silver nanomaterials already in

such products include: metallic silver nanoparticles

(Arora et al.2008; Chi et al.2009; Choi et al.2008;

Hwang et al.2008; Kim et al.2007,2008a,b; Kvitek

et al.2008; Lok et al.2006; Raffi et al.2008; Schrand

et al.2008; Sondi and Salopek-Sondi2004; Vertelov

et al 2008), silver chloride particles (Choi et al

2008), silver-impregnated zeolite powders and

acti-vated carbon materials (Cowan et al 2003; Inoue

et al 2002; Yoon et al 2008a, b), dendrimer–silver

complexes and composites (Balogh et al 2001;

Lesniak et al 2005; Zhang et al 2008),

polymer-silver nanoparticle composites (Bajpai et al 2007;

Damm and Munstedt2008; Damm et al.2008; Hlidek

et al 2008; Jin et al 2007; Kim et al 2009a, b;

Kvitek et al 2008; Naidu et al 2008; Nita 2008;

Sambhy and Sen 2008; Sanpui et al.2008; Xu et al

2006), silver-titanium dioxide composite

nanopow-ders (Yeo and Kang 2008), and silver nanoparticles

coated onto polymers like polyurethane (Jain and

Pradeep 2005) While all of these forms of silver

exert antimicrobial activity to some extent through

release of silver ions, silver nanoparticles might

exhibit additional antimicrobial capabilities not

exerted by bulk or ionic silver (Chen and Schluesener

2008)

Already, silver nanoparticles have been shown to

be effective biocides against: (a) bacteria such as

Escherichia coli, Staphylococcus aureus,

Staphylo-coccus epidermis, Leuconostoc mesenteroides,

Bacil-lus subtilis, Klebsiella mobilis, and Klebsiella

pneumonia among others (Benn and Westerhoff

2008; Chen and Chiang 2008; Falletta et al 2008;

Hernandez-Sierra et al.2008; Ingle et al.2008; Jung

et al 2009; Kim 2007; Kim et al 2007, 2009a, b;

Kvitek et al.2008; Raffi et al.2008; Ruparelia et al

2008; Smetana et al.2008; Sondi and Salopek-Sondi

2004; Vertelov et al 2008; Yang et al 2009; Yoon

et al.2008a,b); (b) fungi such as Aspergillus niger,

Candida albicans, Saccharomyces cerevisia,

Tricho-phyton mentagrophytes, and Penicillium citrinum

(Kim et al 2007, 2008a, b, 2009a, b; Roe et al

2008; Vertelov et al.2008; Zhang et al.2008); and (c)

virii such as Hepatitis B, HIV-1, syncytial virus

(Elechiguerra et al.2005; Lu et al 2008; Sun et al

2008; Zodrow et al 2009) Hybrid silver

nanocom-posites with dendrimers and polymers have been

shown effective for S aureus, Pseudomonas ginosa, E coli, B subtilis, and K mobilis (Balogh

aeru-et al 2001; Zhang et al 2008) Furthermore, silverloaded in nanoporous materials such as silver-exchanged zeolites exhibit antibacterial effects forPseudomonas putida, E coli, B subtilis, S aureus,and P aeruginosa (Cowan et al 2003; Inoue et al

2002; Lind et al.2009; McDonnell et al.2005).Despite the vast number of papers touting thebeneficial antimicrobial effects of silver nanomateri-als, a relatively modest number of studies haveattempted to elucidate the mechanisms by whichsilver nanomaterials exert this antimicrobial activity

As a result, the mechanisms are not widely stood or agreed upon For bacteria, commonlyproposed mechanisms in the literature begin withthe release of silver ions (Hwang et al.2008; Smetana

under-et al 2008) followed by generation of reactiveoxygen species (ROS) (Hwang et al 2008; Kim

et al 2007) and cell membrane damage (Choi et al

2008; Raffi et al 2008; Smetana et al 2008; Sondiand Salopek-Sondi 2004), but there are manycontradictory findings reported

The more widespread our use of silver terials becomes the more widespread will become thepotential for human and ecosystem exposure Silvernanoparticles may be released to the environmentfrom discharges at the point of production, fromerosion of engineered materials in household prod-ucts (e.g., antibacterial coatings and silver-impreg-nated water filters), and from washing or disposal ofsilver-containing products (Benn and Westerhoff

nanoma-2008) Silver released to both natural and engineeredsystems will likely impact the lowest trophic levelsfirst, i.e., bacteria However, little is known abouttrophic transfer of silver and impacts to higherorganisms Indeed, silver nanoparticles have alreadybeen proven toxic to both aerobic and anaerobicbacteria isolated from wastewater treatment plants(Choi and Hu 2008), which we speculate could lead

to severe disruption of this critical environmentalinfrastructure if the load of silver into wastewatertreatment plants increases significantly

Given the vast number of products leveraging thebenefits of silver, it seems prudent to assess thepotential human and ecosystem hazards associatedwith its increased utilization The main routes ofhuman exposure would be the respiratory system,gastrointestinal system, and skin, which are interfaces

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between the internal systems of the human body and

the external environment (Chen and Schluesener

2008) For example, silver nanomaterials may enter

through the respiratory tract due to inhalation of dust

or fumes containing silver nanomaterials at the point

of manufacture, it may be ingested from water,

children’s toys, or food containers treated with silver,

or it may penetrate the skin via silver-containing

textiles and cosmetics Additionally, other potential

entryways could include the female genital tract (due

to incorporation of silver nanoparticles into numerous

female hygienic products), via systemic

administra-tion as it is used for some imaging and therapeutic

purposes (Chen and Schluesener2008; Schrand et al

2008; West and Halas2003), or by incorporation into

medical implants, catheters, and wound dressings

(Furno et al 2004; Galiano et al 2008; Maneerung

et al.2008; Roe et al.2008)

In addition to broad-spectrum antimicrobial

effects, silver nanoparticles have produced toxic

effects in higher cell lines like zebra fish, clams,

rats, and humans (Arora et al.2008, 2009; Asharani

et al 2008; Braydich-Stolle et al 2005; Hsin et al

2008; Hussain et al.2005; Kim et al.2008a,b; Sung

et al 2008; Yeo and Kang 2008; Yeo and Yoon

2009) Evidence in rodents shows that after entering

into the body silver nanoparticles can accumulate

and, in some cases, damage tissues such as the liver,

lungs, and olfactory bulbs, or penetrate the blood–

brain barrier (Arora et al.2009; Braydich-Stolle et al

2005; Hussain et al.2005; Sung et al.2008) A study

in human cells concluded that silver can be genotoxic

(Asharani et al.2009) Additionally, a release of ionic

silver led to the sterility of Macoma balthica clams in

the San Francisco Bay during the 1980s (Brown et al

2003) If silver nanomaterials exhibit similar or

stronger reactivity, the impacts of this isolated event

in San Francisco may foreshadow potential

ecosys-tem impacts of silver nanomaterials

The various forms of silver nanomaterials are

among the most promising antimicrobial agents being

developed from nanotechnology, but the preliminary

evidence of effects on higher organisms alerts us to

remain cautious of its widespread utilization This

cautiousness demands additional research to

deter-mine how to safely design, use, and dispose products

containing silver nanomaterials without creating new

risks to humans or the environment Consequently,

the goal of this article is to provide a critical review

of the state-of-knowledge about silver nanomaterialantibacterial effects with insights toward betterunderstanding potential implications for humanhealth and the environment

Typical forms of silver nanomaterialsHerein, the term ‘‘silver nanomaterials’’ refers to anysilver-containing materials with enhanced activitydue to their nanoscale features In some cases,commercial products containing metallic silver nano-particles in the range of 5–50 nm or ionic silver aregiven the name ‘nanosilver’ (Panyala et al 2008).Silver nanoparticles are nanoscale clusters of metallicsilver atoms, Ag0, engineered for some practicalpurpose—most typically antimicrobial and sterileapplications

The most common method of producing silvernanoparticles is chemical reduction of a silver saltdissolved in water with a reducing compound such asNaBH4, citrate, glucose, hydrazine, and ascorbate(Gulrajani et al.2008; Martinez-Castanon et al.2009;Panacek et al.2006; Pillai and Kamat 2004) Strongreductants lead to small monodisperse particles,while generating larger sizes can be difficult tocontrol Weaker reductants produce slower reductionreactions, but the nanoparticles obtained tend to bemore polydisperse in size In order to generate silvernanoparticles with controlled sizes, a two-stepmethod is usually utilized In this method, nucleiparticles are prepared using a strong reducing agentand they are enlarged by a weak reducing agent(Schneider et al.1994; Shirtcliffe et al.1999) Sincereducing agents for silver nanoparticle synthesis areoften considered toxic or hazardous, the use of greensynthesis methods is becoming a priority (Panacek

et al 2006) A recent review of green synthesismethods for silver nanoparticles discussed the use ofpolysaccharides, polyphenols, Tollens agent, irradia-tion, biological reduction, and polyoxometalate(Sharma et al 2009)

Polysaccharides and polyphenols are typicallyused as capping agents during silver nanoparticlesynthesis, but they may also contribute to reduction

of silver ions through as yet poorly understoodmechanisms For polysaccharides, the reduction ofsilver may be linked to the oxidation of aldehydegroups to carboxylic acid groups (Manzi and van

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Halbeek 1999) Typical polysaccharides used are

glucose, starch, and heparin (Batabyal et al 2007;

Huang and Yang 2004; Manno et al 2008; Singh

et al 2009; Venediktov and Padokhin2008) In the

Tollens method, a silver ammoniacal solution is

reduced by an aldehyde forming silver nanoparticles

This method can be altered (i.e., ‘‘modified Tollens

method’’) by reducing Ag?using saccharides in the

presence of ammonia resulting in films with

nano-particles sizes ranging from 50 to 200 nm and silver

hydrosols ranging from 20 to 50 nm (Panacek et al

2006; Saito et al 2003; Yu and Yam 2004) Silver

nanoparticles can also be synthesized by irradiating

silver salts solutions containing reducing and capping

agents Different sources of irradiation have been

used such as laser, microwave, ionization radiation,

and radiolysis (Abid et al 2002; Ha et al.2006; Li

et al.2006; Long et al.2007; Mahapatra et al.2007;

Pillai and Kamat 2004; Sharma et al 2007, 2008;

Yanagihara et al 2001; Yin et al.2004; Zeng et al

2007)

Biological methods involve the production of

silver nanoparticles utilizing extracts from

bio-organ-isms as reductant, capping agents or both (Li et al

2007; Sanghi and Verma 2009) Such extracts can

include proteins, amino acids, polysaccharides, and

vitamins (Eby et al.2009; Sharma et al.2009) Plant

extracts such as apiin (a glucoside compound) and

leaf extract from magnolia, Persimmon, geranium,

and Pine leaf have also been used as reducing agents

of Ag? to produce silver nanoparticles (Kasthuri

et al.2009; Shankar et al.2003; Song and Kim2009)

Additionally, silver nanoparticles can be synthesized

by several microorganisms such as the bacterial

strains Bacillus licheniformis, K pneumonia, and

fungi strains such as Verticillium and Fusarium

oxysporum, Aspergillus flavus (Ahmad et al 2003;

Kalishwaralal et al 2008; Mokhtari et al 2009;

Mukherjee et al 2001; Senapati et al 2004;

Vig-neshwaran et al 2007) It is not clear if these

microbes are impacted (favorably or unfavorably) by

exposure to engineered forms of nano-scaled silver,

but their seemingly favorable interactions with silver

suggest resistance may be fairly widespread

Another procedure utilized to synthesize silver

nanoparticles is the solvated metal atom dispersion

(SMAD) method (Stoeva et al.2002) In this method,

a metal is co-vaporized with a solvent onto a liquid

nitrogen cooled surface, as liquid nitrogen is removed

the metal atoms and solvent warms causing theaggregation of metal atoms SMAD can be performed

in conjunction with digestive ripening, in this way thenanoparticles resulting from SMAD method arefurther refined by heating them in inert atmosphere

in the presence of selected ligands that encourage theparticles to reach a narrow size range As a result,monodisperse spherical particles are obtained (Sme-tana et al.2005,2008)

The use of silver ions as antimicrobial agents islimited by the solubility of silver ions in biologicaland environmental media containing Cl-, becauseAgCl has a very low solubility and rapidly precip-itates out of solution In some cases, silver salts arestabilized with hyperbranched polymers or dendri-mers that act as nanoreactors, wherein silver ionsare initially complexed with a specific moiety in thepolymeric structure and then reduced to form silvernanoparticles within the polymeric matrix (Fig.1)(Lesniak et al 2005; Zhang et al 2008) Dendri-mer–silver complexes prevent silver ions fromprecipitating and keep silver dispersed in the medialong enough to be delivered where it is desired(Balogh et al 2001; Lesniak et al 2005; Zhang

et al 2008) Silver ions can also be stabilized inzeolite channels (Fig 2) or deposited in activatedcarbon fibers (Inoue et al 2002; Ogden et al 1999;Pal et al 2009)

Composites of silver coatings over titanium ide nanoparticles are used in products such as babybottles and blood-clotting agents to produce antibac-terial activity (Yeo and Kang 2008) Other hybridsilver nanomaterials may include silver nanoparticlescoated onto polyurethane and silver–magnetite com-posite nanoparticles (Fe3O4@Ag); both of thesehybrids are utilized for water disinfection (Gong

diox-et al 2007; Jain and Pradeep 2005) One of thechallenges of using silver (or any) nanoparticles forwater treatment is recovering the particles after thetreatment process Silver–magnetite nanoparticlesoffer the potential advantage of being removed by amagnet, avoiding release to the environment, andmaking possible direct reuse without additionalseparation processes For example, in relatedresearch, magnetite particles proved effective forremoval of arsenic from water (Mayo et al 2007;Yavuz et al.2006) However, for silver materials, anadditional concern is controlling the release of metalions into the final produce water

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Evidence of silver toxicity in microbes

and higher organisms

Here, toxicity refers to any deleterious effects on an

organism upon exposure to silver Obviously, if the

practical intent is to disinfect or sterilize a specific

type of organism, then toxicity may be interpreted

as a positive outcome (e.g., antibacterial, antiviral,

etc.) However, if the same material exerts

unin-tended or undesired impacts to other organisms, then

such toxicity may be interpreted as a potential

hazard

Evidence of toxicity to bacteriaTable 1 presents a concise summary of silver nano-material antibacterial studies Kim et al reported that13.4-nm silver nanoparticles prepared by reduction ofsilver nitrate with sodium borohydride show mini-mum inhibitory concentration (MIC) against E colibelow 6.6 nM and above 33 nM for S aureus (Kim

et al 2007) In another study, 16-nm silver ticles generated by gas condensation were able tocompletely inhibit colony forming units (CFU)ability of E coli at 60 lg/mL (Raffi et al 2008)

nanopar-Fig 1 General formation

Terminal groups on the

surface are marked as

–CH2CH2–CO–Z Silver

ions (represented by M?)

can be pre-organize and

subsequently contained in

the form of solubilized and

stabilize, high-surface area

silver domains Redrawn

based on (Balogh et al.

2001 )

Fig 2 Silver ions

stabilized in zeolite

channels and ionic

exchange of silver ions with

other cations in the media.

Left picture shows zeolite

type A and right picture

shows zeolite type X.

Adapted from (Auerbach

2003 )

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Commercially available silver nanopowders at a

concentration of 300 lg/mL and SMAD-produced

silver nanoparticles at a concentration of 30 lg/mL

were able to reduce (after 10 min contact time)

colony forming units of E coli and S aureus from

2 9 104CFU/mL to 0 and \20, respectively (Smetana

et al.2008)

Additionally, MICs ranging from 13.5 to 1.69 lg/mL

were reported for bacterial strains such as S aureus

CCM 3953, Enterococcus faecalis CCM 4224, E coli

CCM 3954, and P aeruginosa CCM 3955, and for

strains isolated from human clinical material like

P aeruginosa, methicillin-susceptible S epidermidis,

methicillin-resistant S epidermidis, methicillin-resistant

S aureus, vancomycin-resistant Enterococcusfaecium, and K pneumonia when exposed to26-nm silver nanoparticles prepared by the reduc-tion of [Ag(NH3)2]? with D-maltose (Kvitek et al

2008)

Silver nanoparticles also produced 76% reduction

of B subtilis CFU after applying silver nanoparticleswith a size distribution from 14 to 710 nm in anaerosol form (Yoon et al.2008a,b) Likewise, silvernanoparticles are effective against E coli, S aureus,and L mesenteroides Also, 10-nm MyramistinÒstabilized silver nanoparticles inhibit growth of

E coli and S aureus at 2.5 lg/mL and L teroides at 5 lg/mL (Vertelov et al.2008)

mesen-Table 1 Bactericidal activity of nano-scaled silver and silver loaded in zeolite

Silver form Size data Bacterial strain Key aspects References

Silver nanoparticles/

nano-sized silver

powders

13.4 nm b E coli, S aureus Minimal inhibition concentration against

E coli was lower than 6.6 nM and higher than 33 nM for S aureus

Minimal inhibition concentration from 1.69 to 13.5 lg/mL

14.1–710 nmc B subtilis 76% CFU reduction by applying silver

nanoparticles aerosol on B subtilis aerosol

(Yoon et al 2008a ,

b ) Silver nanoparticles

or chloride ions

(Balogh et al 2001 )

Silver zeolite E coli CFU reduced by 7 log units in 5 min (Inoue et al 2002 ) Zeolite containing

silver and zinc

E coli Minimal bactericidal concentration of

78 lg/mL (as Ag?) for bacteria grown is Luria- Bertani broth

Note: Size measured byadynamic light scattering,bTEM images, andcscanning mobility particle sizer

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Dendrimer–silver nanocomposites have also been

proven effective antibacterials For example,

poly(amidoamine) dendrimer–silver composites have

been used against S aureus, P aeruginosa, E coli,

B subtilis, and K mobilis (Balogh et al.2001; Zhang

et al.2008) Additionally, silver ions loaded in zeolites

elicit antibacterial properties Two recent studies

demonstrated 7 log reduction in CFUs for E coli,

from an initial concentration of 107CFU/cm3, after

5 min of contact time with 333.3 lg/mL of Ag-loaded

zeolite (Inoue et al.2002), and MICs of 78 lgAg?/mLfor E coli and 39 lgAg?/mL for S aureus and

P aeruginosa, plus some bactericidal activity againstListeria monocytogenes (Cowan et al.2003)

Evidence of toxicity to other microorganismsSilver nanoparticles also inactivate fungi, virii, andalgae (Table2) For example, silver nanoparticles ofsizes ranging from 1.4 to 7.1 nm and stabilized in

Table 2 Summary of silver nanoparticles toxicity to other microorganisms

Strain Silver nanoparticles Size (nm) Key aspects Reference

1.4–7.1 b Formation of inhibition zones

around silver nanoparticles inoculated spots in agar plates

(Zhang et al 2008 )

S cerevisiae MyramistinÒstabilized

silver nanoparticles

10 a MIC were found to be 5 mg/L (Vertelov et al 2008 )

T mentagrophytes Silver nanoparticles 3 b IC80between 1 and 4 mg/L (Kim et al 2008a , b )

C Albanicas Silver nanoparticles 3 b Silver nanoparticles inhibited

micelial formation, which is responsible for pathogenicity

(Kim et al 2008a , b )

Silver nanoparticles 3b Antifungal activity may be exerted

by cell membrane structure disruption and inhibition of normal budding process

(Kim et al 2009a , b )

Silver nanoparticles coated on plastic catheters

3–18b Catheter coated with silver

nanoparticles inhibited growth and biofilm formation.

1.4–7.1b Formation of inhibition zones

around silver nanoparticles inoculated spots in agar plates

(Zhang et al 2008 )

Viruses

Hepatitis B virus Silver nanoparticles 10b Inhibition of virus replication (Lu et al 2008 )

HIV-1 Silver nanoparticles 16.19 ± 8.68b Only 1–10 nm nanoparticles

attached to virus restraining virus from attaching to host cells.

(Elechiguerra et al 2005 )

Syncitial virus Silver nanoparticles 44% inhibition of Syncitial virus

infection

(Sun et al 2008 ) Algae

C reinhardtii Silver nanoparticles 10–200a EC50for the photosynthetic yield

was found in 0.35 mg/L of total silver content after 1 h of exposure

(Navarro et al 2008 )

Note: Size measured byadynamic light scattering andbTEM images

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hyperbranched polymers, and silver nanoparticles

stabilized with MyramistinÒ (size 10 nm) inhibited

the growth of A niger (Tomsic et al.2009; Vertelov

et al.2008; Zhang et al.2008); the same MyramistinÒ

stabilized particles were found toxic toward S

cere-visiae showing a MIC of 5 mg/L Further, 3-nm silver

nanoparticles showed IC80values from 1 to 4 lg/mL

against T mentagrophytes and 2 to 4 lg/mL for

C albanicans (Kim et al.2008a,b); while in other study,

it was reported that the antifungal activity of silver

nanoparticles against C albanicans could be exerted

by cell membrane structure disruption leading to

reproduction inhibition (Kim et al 2009a, b)

Addi-tionally, silver nanoparticles (sizes ranging from 3 to

18 nm) coated in catheters were able to inhibit growth

of C albicans Although, in this case, no analysis of

the antifungal molecular mechanism was done, the

authors speculate that silver ions (Ag?) released from

the matrix were the antifungal agents (Kim et al

2008a, b, 2009a, b; Roe et al 2008) Furthermore,

silver nanoparticles suppress yeast growth and show

MIC between 6.6 and 13.2 nM (Kim et al 2007)

Silver nanoparticles of sizes from 1.4 to 7.1 nm and

stabilized in hyperbranched polymers

(HPAMAM-N(CH3)2/AgNPs composite) inhibit P citrinum

growth (Zhang et al 2008) Although information

about toxicity for algae is limited, silver nanoparticles

reduced the photosynthetic yield of Chlamydomonas

reinhardtii; in this case, the observed toxicity was

attributed to Ag?ions (Navarro et al.2008)

Evidence of virus inactivation is also reported in

literature For example, silver nanoparticles of 10 nm

are able to inhibit hepatitis B virus replication (Lu

et al 2008) Additionally, PVP-coated silver particles in the range 1–10 nm attach to HIV-1 virus,inhibiting the virus from attaching to host cells(Elechiguerra et al.2005) In other study, PVP-coatedsilver nanoparticles reduced respiratory syncytialvirus infection by 44% (Sun et al.2008) Polysulfoneultrafiltration membranes impregnated with silvernanoparticles of sizes ranging from 1 to 70 nmshowed enhanced virus removal, thus improvingwater disinfection via low pressure membrane filtra-tion (Zodrow et al 2009)

nano-Evidence of toxicity for mammalian cellsSignificant evidence has been reported in relation tothe toxicity of silver nanoparticles to higher organ-isms It has been shown toxic to fish such as zebrafish(Asharani et al.2008; Yeo and Kang2008; Yeo andYoon 2009), Diptera species such as Drosophilamelanogaster, known asfruit fly (Ahamed et al.2010)and different mammalian cell lines of mice (Brayd-ich-Stolle et al.2005; Hussain et al.2005), rats (Kim

et al 2008a, b; Sung et al 2008), and also humans(Asharani et al 2009; Braydich-Stolle et al 2005;Hsin et al 2008; Hussain et al 2005) This reviewpresents only a few, brief examples of silver nano-material toxicity for mammalian cells (Table 3).More detailed, focused reviews on this topic areavailable elsewhere (Chen and Schluesener 2008;Panyala et al.2008)

Table 3 Evidence of nano-scaled silver toxicity for mammalian cells

Rat lung cells Reduction in lung function and inflammatory lesions (Sung et al 2008 )

Sprague-Dawley rats Silver nanoparticles accumulation in olfactory bulb

and subsequent translocation to the brain

(Kim et al 2008a , b ) Mouse stem cells Cell leakage and reduction of mitochondrial function (Braydich-Stolle et al 2005 ) Rat liver cells Cell leakage and reduction of mitochondrial function (Hussain et al 2005 ) Human fibrosarcoma

and human skin/carcinoma

Oxidative stress Low doses produced apoptosis and higher dose necrosis

(Arora et al 2008 ) Mouse fibroblast 50 lg/mL induced apoptosis to 43.4% of cells (Arora et al 2009 )

Human colon cancer 100 lg/mL produced necrosis to 40.2% of cells

Human glioblastoma Silver nanoparticles were found cytotoxic,

genotoxic and antiproliferative

(Asharani et al 2009 ) Human fibroblast Silver nanoparticles were found cytotoxic,

genotoxic and antiproliferative

(Asharani et al 2009 )

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Evidence of silver nanoparticle toxicity for

mam-malian cells was presented in the in vivo studies

performed by Sung et al (2008) and Kim et al

(2008a,b) In the former, a 90-day inhalation study in

rats showed that silver nanoparticles reduce lung

function and produce inflammatory lesions in the

lungs In the later, silver nanoparticles accumulated

in the olfactory bulbs of Sprague-Dawley rats and

also accumulated in the brain

Evidence from in vitro studies is also available in the

literature For example, silver nanoparticles have been

shown to reduce mitochondrial function and to

increase membrane leakage of mouse spermatogonial

stem cell and rat liver cells (Braydich-Stolle et al.2005;

Hussain et al 2005) Studies performed on human

fibrosarcoma and human skin/carcinoma cells with

silver nanoparticles used in a topical antimicrobial

agent concluded that in the presence of the

nanopar-ticles the cellular levels of glutathione are reduced,

indicating oxidative stress, that results in cellular

damage and lipid peroxidation (Arora et al.2008)

However, in the study performed by Arora et al the

dose required to induce apoptosis (0.78–1.56 lg/mL)

was much smaller than that required to produce

necrosis (12.5 lg/mL) in both cell types Therefore,

the authors concluded that, after the required in vivo

studies, it would be possible to define a safe range for

the application of silver nanoparticles as a topical

antimicrobial agent Similar differences of the required

concentration to cause apoptosis or necrosis were

found in a second study by the same authors in mouse

fibroblasts and liver cells (Arora et al.2009) In this

second article, it was suggested that although silver

nanoparticles may enter into the cells, the cellular

antioxidant mechanisms would limit oxidative stress

Mechanistic studies of silver nanoparticle toxicity

in mammalian cells have considered mouse fibroblast

and human colon cancer cells (Hsin et al 2008) In

this study, silver nanoparticle doses of 50 lg/mL

induced apoptosis to 43.4% of fibroblast cells, while

100 lg/mL produced necrosis to 40.2% of the cancer

cells The authors concluded that the apoptotic

mechanisms in fibrosblast cells are a mitochondrial

mediated pathway including the generation of ROS in

the cell, which activate the apoptosis regulators JNK

and p53 proteins inducing protein Bax to migrate to

the surface of the mitochondria That subsequently

induces cytochrome C release from mitochondria and

cleavage of PARP Additionally, in a study done by

Asharani, a possible mechanism of toxicity to humancells was proposed (Asharani et al 2009) Silvernanoparticles would affect the mitochondrial respira-tory chain, causing ROS generation and affecting theproduction of ATP, which subsequently leads to DNAdamage In this study, the authors also concluded that

‘‘even and small dose of Ag-NP (silver nanoparticles)has the potential to cause toxicity’’ and that silvernanoparticles are cytotoxic, genotoxic, and antipro-liferative, being as toxic for human glioblastoma asfor normal human fibroblasts cells

Mechanisms of silver’s antibacterial propertiesAlthough the mechanisms behind the activity ofnano-scaled silver on bacteria are not yet fullyelucidated, the three most common mechanisms oftoxicity proposed to date are: (1) uptake of free silverions followed by disruption of ATP production andDNA replication, (2) silver nanoparticle and silverion generation of ROS, and (3) silver nanoparticledirect damage to cell membranes The variousobserved and hypothesized interactions betweensilver nanomaterials and bacteria cells are conceptu-ally illustrated in Fig.3

Fig 3 Diagram summarizing nano-scaled silver interaction with bacterial cells Nano-scaled silver may (1) release silver ions and generate ROS; (2) interact with membrane proteins affecting their correct function; (3) accumulate in the cell membrane affecting membrane permeability; and (4) enter into the cell where it can generate ROS, release silver ions, and affect DNA Generated ROS may also affect DNA, cell membrane, and membrane proteins, and silver ion release will likely affect DNA and membrane proteins Similar pictures have been published in (Damm et al 2008 ; Neal 2008 )

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Free silver ion uptake

Silver nanoparticles have been reported to dissolve

generating silver ions and it is thought that in vivo this

release would be product of reactions of silver

nano-particles with H2O2(Asharani et al.2009) Asharani has

proposed the following reaction as a possible

mecha-nism for silver nanoparticle oxidative dissolution

2Agþ H2O2þ 2Hþ! 2Agþþ 2H2O E0¼ 0:17 V:

Asharani suggests that in eukaryotic cells this

reaction could occur in the mitochondria, where

exists an important concentration of H? Similarly,

we hypothesized that a similar mechanism could

occur in the bacterial cell membrane where proton

motive force takes place

Another possible mechanism for the oxidative

dissolution of silver nanoparticles has been reported

by Choi et al., in this case silver is oxidized in the

presence of oxygen Choi et al speculated that the

observed changed of color of their silver

nanoparti-cles suspensions, over a week period, would be

attributed to this mechanism

4Agþ O2þ 2H2O$ 4Agþþ 4OH:

The amount of free silver ion measured in this case

was approximately 2.2% of the total silver content in

the silver nanoparticle suspension (Choi et al.2008)

In other article, a 0.1% content of the total silver in

partially oxidized silver nanoparticles suspensions

was attributed to silver ions (Lok et al.2007)

Ionic silver has known antibacterial properties;

thus, it is expected that eluted ions from silver

nanoparticles are responsible for at least a part of

their antibacterial properties At sub-micromolar

concentrations, Ag? interacts with enzymes of the

respiratory chain reaction such as NADH

dehydro-genase resulting in the uncoupling of respiration from

ATP synthesis Silver ions also bind with transport

proteins leading to proton leakage, inducing collapse

of the proton motive force (Dibrov et al.2002; Holt

and Bard 2005; Lok et al.2006) Silver inhibits the

uptake of phosphate and causes the efflux of

intra-cellular phosphate (Schreurs and Rosenberg 1982)

The interaction with respiratory and transport

pro-teins is due to the high affinity of Ag? with thiol

groups present in the cysteine residues of those

proteins (Holt and Bard 2005; Liau et al 1997;Petering1976) Additionally, it has been reported that

Ag? increases DNA mutation frequencies duringpolymerase chain reactions (Yang et al.2009).Bacterial cells exposed to milli-molar Ag? dosessuffer morphological changes such as cytoplasm shrink-age and detachment of cell wall membrane, DNAcondensation and localization in an electron-light region

in the center of the cell, and cell membrane degradationallowing leakage of intracellular contents (Feng et al

2000; Jung et al.2008) Physiological changes occurtogether with the morphological changes Bacterial cellsenter an active, but non-culturable state in whichphysiological levels can be measured but bacteria arenot able to growth and replicate

Several studies have linked the toxicity of silvernanoparticles to the release of silver ions For example,Smetana et al observed that silver ions eroded fromhigh-surface area silver powders prepared by SMADmethod interacted and destroyed bacterial cells (Sme-tana et al 2008) In the same study, a secondpreparation of silver nanoparticles using water-solubleligands was used to obtain silver nanoparticles withhigher surface area to improve their antibacterialefficacy However, the second preparation of silvernanoparticles showed lower toxicity toward bacterialcells than uncoated powders, suggesting the ligandsprevented silver ion erosion; thus, diminishing theresulting toxicity Another possible reason for thisresult is that the surface coatings prevented adhesion

of silver nanoparticles to the bacterial cell surface, butthe authors did not explore this option

Hwang et al observed that Ag?induced the sameeffect in bioluminescence bacteria sensitive to mem-brane protein damage and slightly less effect in a strainsensitive to superoxides compared to silver nanopar-ticles (Hwang et al.2008) The authors suggested thatsilver nanoparticles produce silver ions that moveinside the cell producing ROS through redox reactionswith oxygen In other research, bacterial activity ofactivated carbon fiber supported silver was attributed

to the synergistic action of silver ions, superoxides,and hydrogen peroxide (Le Pape et al.2004).Generation of reactive oxygen speciesReactive oxygen species (ROS) are natural byprod-ucts of the metabolism of respiring organisms While

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