Bioactive constituents with antibacterial, resistance modulation, anti-biofilm formation and efflux pump inhibition properties from Aidia genipiflora stem bark

Abstract Background Antimicrobial resistance is a global health challenge. The involvement of bacterial biofilms and efflux pumps in the development of multidrug resistance (MDR) is well established. Medicinal plants have been proposed as alternatives for combating MDR focusing on their bioactive constituents with resistance modulatory activities. This study was aimed at investigating the stem bark of Aidia genipiflora for bioactive constituents with anti-biofilm, efflux pump inhibition and resistance modulatory activities. Method The crude methanol extract was purified by column chromatography and isolated compounds characterized by mass and nuclear magnetic resonance spectrometry. Antibacterial activity was determined by the High-throughput spot culture growth inhibition and the broth micro-dilution assay. The ethidium bromide accumulation assay was used to determine efflux pump inhibition property. Biofilm inhibition was determined in a microplate crystal violet retention assay. Results Purification of the ethyl acetate fraction led to the isolation of oleanonic acid (1), 4-hydroxy cinnamic acid docosyl ester (2), β-stigmasterol/β-sitosterol (mixture 3a/b) and D-mannitol (4). The minimum inhibitory concentrations (MICs) ranged from 250 to > 500 μg/mL for extracts and fractions and from 15 to 250 μg/mL for compounds. In the presence of sub-inhibitory concentrations of the compounds, the MIC of amoxicillin against E. coli (20 μg/mL) and P. aeruginosa (320 μg/mL) was reduced by 32 and 10 folds respectively. The whole extract demonstrated anti-biofilm formation and efflux pump inhibition in E. coli, S. aureus and P. aeruginosa. The sterol mixture (3a/b) at concentration of 100 μg/mL caused the highest inhibition (73%) of biofilm formation in S. aureus. Oleanonic acid (1) demonstrated remarkable efflux pump inhibition at MIC of 7.8 μg/mL in E. coli better than the standard drugs verapamil and chlorpromazine. Conclusion This study confirms the prospects of A. genipiflora as a source of new antibacterial agents and adjuvants that could interact with some resistance mechanisms in bacteria to enhance the activity of hitherto ineffective antibiotics. “A small portion of the study has been presented in a conference in the form of poster”.

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Antibiotics Application Strategies to Control Biofilm Formation in Pathogenic Bacteria

Current Pharmaceutical Biotechnology ◽

10.2174/1389201020666191112155905 ◽

2020 ◽

Vol 21 (4) ◽

pp. 270-286 ◽

Cited By ~ 2

Author(s):

Fazlurrahman Khan ◽

Dung T.N. Pham ◽

Sandra F. Oloketuyi ◽

Young-Mog Kim

Keyword(s):

Biofilm Formation ◽

Pathogenic Bacteria ◽

Extracellular Polymeric Substances ◽

Quorum Quenching ◽

Resistance Mechanisms ◽

Bacterial Biofilm ◽

Bacterial Cells ◽

High Concentration ◽

Biofilm Inhibition ◽

Abiotic Surfaces

Background: The establishment of a biofilm by most pathogenic bacteria has been known as one of the resistance mechanisms against antibiotics. A biofilm is a structural component where the bacterial community adheres to the biotic or abiotic surfaces by the help of Extracellular Polymeric Substances (EPS) produced by bacterial cells. The biofilm matrix possesses the ability to resist several adverse environmental factors, including the effect of antibiotics. Therefore, the resistance of bacterial biofilm-forming cells could be increased up to 1000 times than the planktonic cells, hence requiring a significantly high concentration of antibiotics for treatment. Methods: Up to the present, several methodologies employing antibiotics as an anti-biofilm, antivirulence or quorum quenching agent have been developed for biofilm inhibition and eradication of a pre-formed mature biofilm. Results: Among the anti-biofilm strategies being tested, the sub-minimal inhibitory concentration of several antibiotics either alone or in combination has been shown to inhibit biofilm formation and down-regulate the production of virulence factors. The combinatorial strategies include (1) combination of multiple antibiotics, (2) combination of antibiotics with non-antibiotic agents and (3) loading of antibiotics onto a carrier. Conclusion: The present review paper describes the role of several antibiotics as biofilm inhibitors and also the alternative strategies adopted for applications in eradicating and inhibiting the formation of biofilm by pathogenic bacteria.

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Effect of Trans-Cinnamaldehyde on preformed biofilm and biofilm formation of Escherichia coli isolated from urine specimens

QJM ◽

10.1093/qjmed/hcab101 ◽

2021 ◽

Vol 114 (Supplement_1) ◽

Author(s):

Eman Adel El-Haddad ◽

Soha Abdel Rahman El-Hady ◽

Amira Esmail Abdel Hamid ◽

Hisham Abdel Majeed Fahim

Keyword(s):

Escherichia Coli ◽

Biofilm Formation ◽

Defense Mechanisms ◽

Colorimetric Assay ◽

Biofilm Inhibition ◽

Hospital Acquired Infection ◽

E Coli ◽

Before And After ◽

Tract Infections ◽

Hospital Acquired

Abstract Introduction Bacteria in most environments exist as communities of sessile cells in a selfproduced polymeric matrix known as biofilms. Biofilms are responsible for more than 80% of infections, including urinary tract infections (UTI). UTI is the most common hospital acquired infection, caused mainly by Escherichia coli (E.coli). E. coli can readily form biofilm in such infections, specially in the presence of indwelling urinary catheter. It’s difficult to eradicate bacteria in biofilms, since they are shielded from the host defense mechanisms as phagocytes and antibodies, as well as antibiotics. Searching for alternative or adjuvant substances for prevention and eradication of biofilm associated infections are therefore urgently needed. Aim of the work Studying the efficacy of the trans-cinnamaldehyde (TC) for preventing E. coli biofilm formation. Materials and methods Thirty isolates of E.coli were obtained from urine samples. To test the effect of TC on E.coli biofilm formation and preformed biofilms, microtitre plates (MTP) were inoculated with the isolated E.coli and were treated with different concentrations of TC and incubated at 37° C. A colorimetric assay was used to assess biofilm inhibition and inactivation and optical densities (OD) were compared before and after adding different TC concentrations. Results The mean OD of the isolated E.coli biofilms was 1.3 and significantly decreased when mixed with TC different concentrations. TC had high activity in inhibition of preformed E.coli biofilms, where no biofilm was detected on MTP treated with 1.25% and 1.5% TC. Conclusion TC inhibited the biofilm forming ability of E.coli isolates could fully inactivate formed biofilms, suggesting its possibility to be used as an anti-biofilm agent or adjuvant in preventing and treating UTI caused by biofilm producing E.coli.

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Significance of CSE-1034 (Elores™) in Treatment of Urinary Tract Infections due to Multi Drug Resistant ESBLPositiveEscherichia coli

Asian Journal of Pharmaceutical and Clinical Research ◽

10.22159/ajpcr.2016.v9i6.14409 ◽

2016 ◽

Vol 9 (6) ◽

pp. 12

Author(s):

Vishal Bhambri

Keyword(s):

Biofilm Formation ◽

Geriatric Patients ◽

Efflux Pump ◽

Drug Resistant ◽

Over Expression ◽

E Coli ◽

Disodium Edetate ◽

Tract Infections ◽

Hospital Acquired ◽

Common Infections

UTIs are one of the most common infections of geriatric patients in community acquired and hospital acquired settings. Escherichia coli(E. coli) is usually the most common pathogen responsible for UTI.Antibiotic resistance is the main reason for failure of therapy especially in E. coli. Biofilm formation, ESBL and MBL production&efflux pump over expression are the main reasons for antibiotic resistance in E. coli. That’s why management of UTI with multi drug resistant (MDR)E. coli has always been challenge for the clinicians.We are reporting empiric use of new antibiotic adjuvant entity Elores™(Ceftriaxone/ Sulbactam/Disodium-edetate) in the management of UTIs caused by multi drug resistant E. coli.

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Intensity and Mechanisms of Fluoroquinolone Resistance within theH30 andH30Rx Subclones of Escherichia coli Sequence Type 131 Compared with Other Fluoroquinolone-Resistant E. coli

Antimicrobial Agents and Chemotherapy ◽

10.1128/aac.00673-15 ◽

2015 ◽

Vol 59 (8) ◽

pp. 4471-4480 ◽

Cited By ~ 37

Author(s):

James R. Johnson ◽

Brian Johnston ◽

Michael A. Kuskowski ◽

Evgeni V. Sokurenko ◽

Veronika Tchesnokova

Keyword(s):

Escherichia Coli ◽

Efflux Pump ◽

Resistance Mechanisms ◽

Sequence Type ◽

Fluoroquinolone Resistance ◽

Solvent Tolerance ◽

Organic Solvent Tolerance ◽

Content Type ◽

E Coli ◽

Pump Activity

ABSTRACTThe recent expansion of theH30 subclone ofEscherichia colisequence type 131 (ST131) and its CTX-M-15-associatedH30Rx subset remains unexplained. Although ST131H30 typically exhibits fluoroquinolone resistance, so do multiple otherE. colilineages that have not expanded similarly. To determine whetherH30 isolates have more intense fluoroquinolone resistance than other fluoroquinolone-resistantE. coliisolates and to identify possible mechanisms, we determined the MICs for four fluoroquinolones (ciprofloxacin, levofloxacin, moxifloxacin, and norfloxacin) among 89 well-characterized, genetically diverse fluoroquinolone-resistantE. coliisolates (48 non-H30 and 41H30 [23H30Rx and 18H30 non-Rx]). We compared the MICs with theH30 andH30Rx status, the presence/number of nonsynonymous mutations ingyrA,parC, andparE, the presence ofaac(6′)-1b-cr(an aminoglycoside/fluoroquinolone agent-modifying enzyme), and the efflux pump activity (measured as organic solvent tolerance [OST]). Among 1,518 recentE. coliclinical isolates, ST131H30 predominated clonally, both overall and among the fluoroquinolone-resistant isolates. Among the 89 study isolates, compared with non-H30 isolates,H30 isolates exhibited categorically higher MICs for all four fluoroquinolone agents, higher absolute ciprofloxacin and norfloxacin MICs, more nonsynonymous mutations ingyrA,parC, andparE(specificallygyrAD87N,parCE84V, andparEI529L), and a numerically higher prevalence of (H30Rx-associated)aac(6′)-1b-crbut lower OST scores. All putative resistance mechanisms were significantly associated with the MICs [foraac(6′)-1b-cr: ciprofloxacin and norfloxacin only].parCD87N corresponded with ST131H30 andparEI529L with ST131 generally. Thus, more intense fluoroquinolone resistance may provide ST131H30, especiallyH30Rx [ifaac(6′)-1b-crpositive], with subtle fitness advantages over other fluoroquinolone-resistantE. colistrains. This urges both parsimonious fluoroquinolone use and a search for other fitness-enhancing traits within ST131H30.

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The Inhibition Activity of Tannin on the Formation of Mono-Species and Polymicrobial Biofilm Escherichia coli, Staphylococcus aureus, Pseudomonas aeruginosa, and Candida albicans

Majalah Obat Tradisional ◽

10.22146/mot.44532 ◽

2019 ◽

Vol 24 (2) ◽

pp. 110 ◽

Cited By ~ 1

Author(s):

Hasyrul Hamzah ◽

Triana Hertiani ◽

Sylvia Utami Tunjung Pratiwi ◽

Titik Nuryastuti

Keyword(s):

Biofilm Formation ◽

Inhibitory Concentration ◽

Potential Candidate ◽

Inhibition Activity ◽

Middle Phase ◽

Biofilm Inhibition ◽

E Coli ◽

The Social ◽

The Matrix ◽

Anti Fungal

Biofilm acts as the mediator for infection nowadays. Approximately, more than 80% infection incidents are biofilm-formation related. Biofilm as bacteria's defense system is more difficult to eradicate by antibiotic; therefore, pathogen bacteria on their biofilm forms can make serious problems for human health. The invention of a new candidate for polymicrobial biofilm can be an essential challenge to be studied, in order to prevent infections related to biofilm. Tannin is a polyphenol compound with anti-bacterial and anti-fungal potential. This study aims to acknowledge the effectiveness of tannin in inhibition and degradation of C. albicans, P. aeruginosa, E. coli, S. aureus, and polymicrobial biofilm. The assay for biofilm inhibition and degradation were determined with microtiter broth method. The effectivity of tannin antibiofilm against polymicrobial biofilm were analyzed by calculating minimum biofilm inhibitory concentration (MBIC50) and minimum biofilm eradication concentration (MBEC50) values. The mechanism of action of tannin against polymicrobial biofilm was tested using scanning electron microscopy (SEM). The data were analyzed using the Statistical Package for the Social Sciences (SPSS) with a 95% confidence level. Tannin 1% gave inhibition activity of mono-species biofilm formation S. aureus in the middle phase and maturation of 79.04±0.01, 61.48±0.03, E. coli 74.56±0.01, 67.91±0.02, P. aeruginosa 67.32±0.05, 35.13± 0.01, C. albicans 60.62±0.01, 47.16±0.01. The results also provide evidence that tannin activity can degrade and damage the matrix of extracellular polymeric substance (EPS) polymicrobial biofilms. Hence, tannins can be a potential candidate for new antibiofilm for polymicrobial biofilm.

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Truncation of poly-N-acetylglucosamine (PNAG) polymerization with N-acetylglucosamine analogues

10.1101/2021.10.13.464265 ◽

2021 ◽

Author(s):

Zachary Morrison ◽

Alexander Eddenden ◽

Adithya S Subramanian ◽

P. Lynne Howell ◽

mark nitz

Keyword(s):

Biofilm Formation ◽

Chain Termination ◽

Salvage Pathway ◽

Biofilm Inhibition ◽

E Coli ◽

Substrate Analogues

Bacteria require polysaccharides for structure, survival, and virulence. Despite the central role these structures play in microbiology few tools are available to manipulate their production. In E. coli the glycosyltransferase complex PgaCD produces poly-N-acetylglucosamine (PNAG), an extracellular matrix polysaccharide required for biofilm formation. We report that C6-substituted (H, F, N3, SH, NH2) UDP-GlcNAc substrate analogues are inhibitors of PgaCD. In vitro the inhibitors cause PNAG chain termination; consistent with the mechanism of PNAG polymerization from the non-reducing terminus. In vivo, expression of the GlcNAc-1-kinase NahK in E. coli provided a non-native GlcNAc salvage pathway that produced the UDP-GlcNAc analogue inhibitors in situ. The 6-fluoro and 6-deoxy derivatives were potent inhibitors of biofilm formation in the transformed strain, providing a tool to manipulate this key exopolysaccharide. Characterization of the UDP-GlcNAc pool and quantification of PNAG generation support PNAG termination as the primary in vivo mechanism of biofilm inhibition by 6-fluoro UDP-GlcNAc.

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Novel Strategy for Biofilm Inhibition by Using Small Molecules Targeting Molecular Chaperone DnaK

Antimicrobial Agents and Chemotherapy ◽

10.1128/aac.04465-14 ◽

2014 ◽

Vol 59 (1) ◽

pp. 633-641 ◽

Cited By ~ 49

Author(s):

Ken-ichi Arita-Morioka ◽

Kunitoshi Yamanaka ◽

Yoshimitsu Mizunoe ◽

Teru Ogura ◽

Shinya Sugimoto

Keyword(s):

Molecular Chaperone ◽

Biofilm Formation ◽

Antimicrobial Agents ◽

Dependent Manner ◽

Cellular Functions ◽

Biofilm Inhibition ◽

Content Type ◽

E Coli ◽

Chaperone Dnak ◽

Biofilm Development

ABSTRACTBiofilms are complex communities of microorganisms that attach to surfaces and are embedded in a self-produced extracellular matrix. Since these cells acquire increased tolerance against antimicrobial agents and host immune systems, biofilm-associated infectious diseases tend to become chronic. We show here that the molecular chaperone DnaK is important for biofilm formation and that chemical inhibition of DnaK cellular functions is effective in preventing biofilm development. Genetic, microbial, and microscopic analyses revealed that deletion of thednaKgene markedly reduced the production of the extracellular functional amyloid curli, which contributes to the robustness ofEscherichia colibiofilms. We tested the ability of DnaK inhibitors myricetin (Myr), telmisartan, pancuronium bromide, and zafirlukast to prevent biofilm formation ofE. coli. Only Myr, a flavonol widely distributed in plants, inhibited biofilm formation in a concentration-dependent manner (50% inhibitory concentration [IC50] = 46.2 μM); however, it did not affect growth. Transmission electron microscopy demonstrated that Myr inhibited the production of curli. Phenotypic analyses of thermosensitivity, cell division, intracellular level of RNA polymerase sigma factor RpoH, and vulnerability to vancomycin revealed that Myr altered the phenotype ofE. coliwild-type cells to make them resemble those of the isogenicdnaKdeletion mutant, indicating that Myr inhibits cellular functions of DnaK. These findings provide insights into the significance of DnaK in curli-dependent biofilm formation and indicate that DnaK is an ideal target for antibiofilm drugs.

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Phytochemical analysis of medicinal plants of Nepal and their antibacterial and antibiofilm activities against uropathogenic Escherichia coli

BMC Complementary Medicine and Therapies ◽

10.1186/s12906-021-03293-3 ◽

2021 ◽

Vol 21 (1) ◽

Author(s):

Sudip Bhandari ◽

Karan Khadayat ◽

Sami Poudel ◽

Sunil Shrestha ◽

Raju Shrestha ◽

...

Keyword(s):

Escherichia Coli ◽

Medicinal Plants ◽

Biofilm Formation ◽

Prunus Persica ◽

Inhibitory Concentration ◽

Total Phenolic ◽

Antibiofilm Activity ◽

Phytochemical Analysis ◽

Biofilm Inhibition ◽

E Coli

Abstract Background A biofilm is an extracellular polymeric substance (EPS) composed of polysaccharides, proteins, nucleic acids, and lipids that impede antibiotics and immune cells, thus providing a shielded environment for bacterial growth. Due to biofilm formation, some microbes can show up to 1000 fold increased resistance towards the antibiotics than the normal planktonic forms. The study was conducted to screen the crude extracts of medicinal plants used in Nepal for their in vitro antibiofilm activities. Methods Total phenolic and total flavonoid contents were determined by using a Folin-Ciocalteau reagent and aluminium trichloride method, respectively. Resazurin assay was used to determine the minimum inhibitory concentration (MIC) and minimum bactericidal concentration (MBC). The initial antibiofilm activities and their inhibitory concentration (IC50) values were determined by the microtiter based modified crystal violet staining method. Results Out of 25 different plant extracts were used for the study, methanolic extracts of 20 plants showed a biofilm inhibition activity against five different strong biofilm producing Escherichia coli strains. Calotropis gigantea exhibited inhibition against all five different E. coli strains with IC50 values ranging from 299.7 ± 20.5 to 427.4 ± 2.7 μg/mL. Apart from that, Eclipta prostrata also showed biofilm formation inhibition, followed by Eupatorium adenophorum, Moringa oleifera, Ocimum tenuifolium, Oxalis lantifolia, Prunus persica, and Urtica parviflora. The extracts of C. gigantea, E. prostrata, Mangifera indica, O. tenuifolium, P. persica, and U. parviflora exhibited a moderate to poor MIC value ranging from 625 to 2500 μg/mL. The highest amount of phenolic content (TPC) was found in Acacia catechu followed by Morus alba, which was 38.9 and 25.1 mg gallic acid equivalents, respectively. The highest amount of flavonoid content was found in A. catechu followed by M. indica, which was 27.1 and 20.8 mg quercetin equivalents, respectively. Conclusion Extracts of C. gigantea, E. prostrata, P. persica, U. parviflora, and O. tenuifolium showed antibacterial as well as antibiofilm activity against pathogenic and strong biofilm producing E. coli. Thus, extracts or the pure compound from these medicinal plants could be used as antibiotics in the future.

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In vitro inhibition of multi-drug resistant Pseudomonas efflux pump by Xylopia aethiopica (Dunal) A. Rich

AROC in Pharmaceutical and Biotechnology ◽

10.53858/arocpb01022027 ◽

2021 ◽

Vol 1 (2) ◽

pp. 20-27

Author(s):

Jeremiah John Oloche ◽

◽

Bolaji Bosede Oluremi ◽

Temiloluwa Oyindamola Koya

Keyword(s):

Antimicrobial Activity ◽

Ethyl Acetate ◽

Bacterial Infections ◽

Efflux Pump ◽

Ethyl Acetate Fraction ◽

Botanical Garden ◽

Stem Bark ◽

Antibiotics Resistance ◽

Xylopia Aethiopica

Global health is under constant threat due to antimicrobial drug resistance. Bacterial Infections caused by Pseudomonas aeruginosa are of importance because of their antibiotics resistance. This study aimed at evaluating the effects of extracts of Xylopia aethiopica (XA) on multidrug-resistant (MDR) Pseudomonas isolates. Fresh samples of XA leaf, stem bark and roots were collected from the botanical garden, University of Ibadan, Nigeria. Dried and pulverized samples were extracted with methanol and partitioned into n-hexane, dichloromethane and ethyl acetate. Phytochemical screening of the extracts was performed by standard methods. Antimicrobial activity and synergistic interaction were determined using microdilution and checkerboard broth dilution methods, respectively. The results revealed that crude methanol extracts of XA leaf, stem bark and root significantly (p<0.05) inhibited the growth of all tested MDR Pseudomonas isolates at 10 mg/mL. At 1 mg/mL, the ethyl acetate fraction of the leaf, and dichloromethane fraction of the roots produced clear zones of inhibition of 12 – 20 mm, and minimum inhibitory concentrations (MICs) of 1 µg/mL and 0.5 mg/mL, respectively. The modulation factor (MF) of ciprofloxacin, dichloromethane fraction of XA roots and ethyl acetate fraction of XA leaf were 4, 8, and 4 on MDR isolates E01006, OAU058 and P. aeruginosa ATCC 27853, respectively. In all tested isolates, but not E01006 and E01024, the fractional MICs of ciprofloxacin/ethylacetate fraction of XA leaf extract combination was not significantly different (p>0.05) compared with ciprofloxacin/verapamil combination. In conclusion, the root and leaf fractions Xylopia aethiopica that demonstrated antimicrobial activity against MDR P. aeruginosa and synergised with ciprofloxacin have the potential to rejuvenate the antimicrobial activity of ciprofloxacin in MDR P. aeruginosa.

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Effect of Spermidine on Biofilm Formation in Escherichia coli K-12

Journal of Bacteriology ◽

10.1128/jb.00652-20 ◽

2021 ◽

Vol 203 (10) ◽

Author(s):

Kullathida Thongbhubate ◽

Yuko Nakafuji ◽

Rina Matsuoka ◽

Sonomi Kakegawa ◽

Hideyuki Suzuki

Keyword(s):

Escherichia Coli ◽

Biofilm Formation ◽

Efflux Pump ◽

Conclusive Evidence ◽

Bacterial Biofilm ◽

Spermidine Synthase ◽

Periplasmic Binding Protein ◽

Content Type ◽

E Coli ◽

K 12

ABSTRACT Polyamines are essential for biofilm formation in Escherichia coli, but it is still unclear which polyamines are primarily responsible for this phenomenon. To address this issue, we constructed a series of E. coli K-12 strains with mutations in genes required for the synthesis and metabolism of polyamines. Disruption of the spermidine synthase gene (speE) caused a severe defect in biofilm formation. This defect was rescued by the addition of spermidine to the medium but not by putrescine or cadaverine. A multidrug/spermidine efflux pump membrane subunit (MdtJ)-deficient strain was anticipated to accumulate more spermidine and result in enhanced biofilm formation compared to the MdtJ+ strain. However, the mdtJ mutation did not affect intracellular spermidine or biofilm concentrations. E. coli has the spermidine acetyltransferase (SpeG) and glutathionylspermidine synthetase/amidase (Gss) to metabolize intracellular spermidine. Under biofilm-forming conditions, not Gss but SpeG plays a major role in decreasing the too-high intracellular spermidine concentrations. Additionally, PotFGHI can function as a compensatory importer of spermidine when PotABCD is absent under biofilm-forming conditions. Last, we report here that, in addition to intracellular spermidine, the periplasmic binding protein (PotD) of the spermidine preferential ABC transporter is essential for stimulating biofilm formation. IMPORTANCE Previous reports have speculated on the effect of polyamines on bacterial biofilm formation. However, the regulation of biofilm formation by polyamines in Escherichia coli has not yet been assessed. The identification of polyamines that stimulate biofilm formation is important for developing novel therapies for biofilm-forming pathogens. This study sheds light on biofilm regulation in E. coli. Our findings provide conclusive evidence that only spermidine can stimulate biofilm formation in E. coli cells, not putrescine or cadaverine. Last, ΔpotD inhibits biofilm formation even though the spermidine is synthesized inside the cells from putrescine. Since PotD is significant for biofilm formation and there is no ortholog of the PotABCD transporter in humans, PotD could be a target for the development of biofilm inhibitors.

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