In Vitro Oxidative Crosslinking of Recombinant Barnacle Cyprid Cement Gland Proteins

AbstractBarnacle adhesion is a focus for fouling-control technologies as well as the development of bioinspired adhesives, although the mechanisms remain very poorly understood. The barnacle cypris larva is responsible for surface colonisation. Cyprids release cement from paired glands that contain proteins, carbohydrates and lipids, although further compositional details are scant. Several genes coding for cement gland-specific proteins were identified, but only one of these showed database homology. This was a lysyl oxidase-like protein (lcp_LOX). LOX-like enzymes have been previously identified in the proteome of adult barnacle cement secretory tissue. We attempted to produce recombinant LOX in E. coli, in order to identify its role in cyprid cement polymerisation. We also produced two other cement gland proteins (lcp3_36k_3B8 and lcp2_57k_2F5). lcp2_57k_2F5 contained 56 lysine residues and constituted a plausible substrate for LOX. While significant quantities of soluble lcp3_36k_3B8 and lcp2_57k_2F5 were produced in E. coli, production of stably soluble lcp_LOX failed. A commercially sourced human LOX catalysed the crosslinking of lcp2_57k_2F5 into putative dimers and trimers, and this reaction was inhibited by lcp3_36k_3B8. Inhibition of the lcp_LOX:lcp2_57k_2F5 reaction by lcp3_36k_3B8 appeared to be substrate specific, with no inhibitory effect on the oxidation of cadaverine by LOX. The results demonstrate a possible curing mechanism for barnacle cyprid cement and, thus, provide a basis for a more complete understanding of larval adhesion for targeted control of marine biofouling and adhesives for niche applications.

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Flavone-Rich Fractions and Extracts from Oroxylum indicum and Their Antibacterial Activities against Clinically Isolated Zoonotic Bacteria and Free Radical Scavenging Effects

Molecules ◽

10.3390/molecules26061773 ◽

2021 ◽

Vol 26 (6) ◽

pp. 1773

Author(s):

Patchima Sithisarn ◽

Piyanuch Rojsanga ◽

Pongtip Sithisarn

Keyword(s):

Antibacterial Activities ◽

Inhibitory Effect ◽

Total Flavonoid ◽

Total Phenolic ◽

Antibacterial Effects ◽

E Coli ◽

Young Fruit ◽

Oroxylum Indicum ◽

Zoonotic Bacteria

Oroxylum indicum extracts from the seeds collected from Lampang and Pattani provinces in Thailand, and young fruits and flowers exhibited in vitro display antioxidant and antibacterial activities against clinically isolated zoonotic bacteria including Staphylococcus intermedius, Streptococcus suis, Pseudomonas aeruginosa, β-hemolytic Escherichia coli and Staphylococcus aureus. The orange crystals and yellow precipitates were obtained from the preparation processes of the seed extracts. The orange-red crystals from the seeds collected from Lampang province exhibited strong in vitro 2,2-diphenyl-1-picrylhydrazyl (DPPH) scavenging effects (EC50 value = 25.99 ± 3.30 μg/mL) and antibacterial effects on S. intermedius and β-hemolytic E. coli while the yellow precipitate from the same source exhibited only antioxidant activity. Quantitative analysis of phytochemicals in O. indicum samples by spectrophotometric and HPLC techniques showed that they contained different amounts of total phenolic, total flavonoid and three major flavones; baicalin, baicalein and chrysin contents. Young fruit extract, which contained low amounts of flavone contents, still promoted antibacterial effects against the tested bacteria with IC50 values lower than 1 mg/mL and MIC values between 4 to 10 mg/mL in S. intermedius, S. aureus and S suis while higher IC50 and MIC values against P. aeruginosa and β-hemolytic E. coli were found. From scanning electron microscopy, the extract of the young fruit of O. indicum promoted morphological changes in the bacterial cells by disrupting the bacterial cell walls, inducing leakage of the cellular content, and generating the abnormal accumulation of cells. The mechanism of action of the extract for this antibacterial effect may be the disruption of the cell membrane and abnormal cell aggregations. Regression analysis of the results suggests the correlation between total phenolic and total flavonoid contents and antioxidant and antibacterial effects. Baicalin was found to have a high correlation with an inhibitory effect against β-hemolytic E. coli while three unidentified peaks, which could be flavones, showed high correlations with an inhibitory effect against S. intermedius, S. suis, P. aeruginosa and S. aureus.

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Lysyl oxidase activity and elastin/glycosaminoglycan interactions in growing chick and rat aortas.

The Journal of Cell Biology ◽

10.1083/jcb.105.3.1463 ◽

1987 ◽

Vol 105 (3) ◽

pp. 1463-1469 ◽

Cited By ~ 60

Author(s):

C Fornieri ◽

M Baccarani-Contri ◽

D Quaglino ◽

I Pasquali-Ronchetti

Keyword(s):

Hydrophobic Interactions ◽

Isonicotinic Acid ◽

Lysyl Oxidase ◽

Acid Hydrazide ◽

Elastic Fibers ◽

Amino Groups ◽

Lysine Residues ◽

Oxidase Inhibition

Hydrophobic tropoelastin molecules aggregate in vitro in physiological conditions and form fibers very similar to natural ones (Bressan, G. M., I. Pasquali Ronchetti, C. Fornieri, F. Mattioli, I. Castellani, and D. Volpin, 1986, J. Ultrastruct. Molec. Struct. Res., 94:209-216). Similar hydrophobic interactions might be operative in in vivo fibrogenesis. Data are presented suggesting that matrix glycosaminoglycans (GAGs) prevent spontaneous tropoelastin aggregation in vivo, at least up to the deamination of lysine residues on tropoelastin by matrix lysyl oxidase. Lysyl oxidase inhibitors beta-aminopropionitrile, aminoacetonitrile, semicarbazide, and isonicotinic acid hydrazide were given to newborn chicks, to chick embryos, and to newborn rats, and the ultrastructural alterations of the aortic elastic fibers were analyzed and compared with the extent of the enzyme inhibition. When inhibition was greater than 65% all chemicals induced alterations of elastic fibers in the form of lateral aggregates of elastin, which were always permeated by cytochemically and immunologically recognizable GAGs. The number and size of the abnormal elastin/GAGs aggregates were proportional to the extent of lysyl oxidase inhibition. The phenomenon was independent of the animal species. All data suggest that, upon inhibition of lysyl oxidase, matrix GAGs remain among elastin molecules during fibrogenesis by binding to positively charged amino groups on elastin. Newly synthesized and secreted tropoelastin has the highest number of free epsilon amino groups, and, therefore, the highest capability of binding to GAGs. These polyanions, by virtue of their great hydration and dispersing power, could prevent random spontaneous aggregation of hydrophobic tropoelastin in the extracellular space.

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Synthesis of Some Polysubstituted Nicotinonitriles and Derived Pyrido[2,3-d]pyrimidines asIn VitroCytotoxic and Antimicrobial Candidates

Journal of Chemistry ◽

10.1155/2016/2135893 ◽

2016 ◽

Vol 2016 ◽

pp. 1-12 ◽

Cited By ~ 5

Author(s):

Hassan M. Faidallah ◽

Sherif A. F. Rostom ◽

Khalid A. Khan

Keyword(s):

Cell Line ◽

Colon Carcinoma ◽

Human Tumor ◽

Human Colon ◽

Pyrimidine Ring ◽

Inhibitory Effect ◽

Breast Adenocarcinoma ◽

Human Tumor Cell Lines ◽

E Coli

The synthesis of polysubstituted pyridines, in addition to some derived pyrido[2,3-d]pyrimidine ring systems supported with chemotherapeutically active functionalities, is described. They were evaluated for theirin vitrocytotoxic effects against three different human tumor cell lines (human colon carcinoma HT29, hepatocellular carcinoma Hep-G2, and Caucasian breast adenocarcinoma MCF7). Nine compounds displayed variable cytotoxic potential, among which alkylthio analogs33,34, and37emerged as the most active members, being almost twice as active as doxorubicin against the colon carcinoma HT29 cell line. In addition, the same three analogs showed a clear differential cytotoxic profile as they exhibited a marginal inhibitory effect on the growth of the normal nontransformed human foreskin fibroblast Hs27 cell line. Meanwhile, nineteen compounds were able to exhibit significant antibacterial activity against both Gram-positive and Gram-negative bacteria, together with moderate antifungal activities. The pyrido[2,3-d]pyrimidine-2(1H)-thione30together with its alkylthio derivatives33and34stemmed as the most active antimicrobial members being equipotent to ampicillin againstS. aureus,E. coli,andP. aeruginosa,together with a noticeable antifungal activity againstC. albicans.Compounds33and34could be considered as a promising template for possible dual antimicrobial-anticancer candidates.

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Lactobacillus inhibitor production against Escherichia coli and coaggregation ability with uropathogens

Canadian Journal of Microbiology ◽

10.1139/m88-063 ◽

1988 ◽

Vol 34 (3) ◽

pp. 344-351 ◽

Cited By ~ 159

Author(s):

Gregor Reid ◽

Jacqueline A. McGroarty ◽

Rosanne Angotti ◽

Roger L. Cook

Keyword(s):

Escherichia Coli ◽

Defense Mechanism ◽

Inhibitory Effect ◽

Second Phase ◽

Vitro Assay ◽

E Coli ◽

Host Defense Mechanism ◽

Important Host ◽

Two Phases

Previous investigations have shown that certain strains of lactobacilli can competitively exclude uropathogens from attaching to uroepithelial cells and from causing urinary tract infection in animals. The finding of an inhibitory effect produced by Lactobacillus casei ssp. rhamnosus GR-1 against the growth of uropathogens was investigated further using two Escherichia coli indicator strains Hu 734 and ATCC 25922. There were two phases to the inhibitor studies. The first one using an agar sandwich technique showed that the inhibitor activity was heat stable and inhibitory to the E. coli. The second phase showed that MRS broth provided optimum lactobacilli growth and inhibitor production. In addition, the inhibition was present under conditions buffering for acid and pH. The data indicated that the inhibitory effect was not due to bacteriophages or hydrogen peroxide. Strain GR-1 was found to coaggregate with E. coli ATCC 25922 in urine, a phenomenon that has not previously been reported for urogenital bacteria. An in vitro assay system was developed to study the coaggregation of various lactobacilli and uropathogens. The results demonstrated that highest coaggregation scores occurred after 4 h incubation at 37 °C with lactobacilli and two type-1 fimbriated E. coli strains. Of the nine lactobacilli strains tested, each was found to coaggregate with 2 or more of the 13 uropathogens. The dominance of inhibitor-producing lactobacilli on the urogenital epithelium and the ability of these organisms to interact closely with uropathogens would constitute an important host defense mechanism against infection.

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Evaluation of mesoporous silica nanoparticles-based nanoantibiotics and capsaicin on E. coli and S. aureus biofilms

10.5194/biofilms9-133 ◽

2020 ◽

Author(s):

Anna Slita ◽

Prakirth Govardhanam ◽

Ida Opstad ◽

Didem Sen Karaman ◽

Jessica Rosenholm

Keyword(s):

Mesoporous Silica ◽

Silica Nanoparticles ◽

High Throughput ◽

Mesoporous Silica Nanoparticles ◽

Inhibitory Effect ◽

E Coli ◽

Resazurin Assay ◽

Antibiotic Drug ◽

High Throughput Assay

Introduction Since antibiotics were discovered, bacteria have demonstrated the ability to develop resistance by many different mechanisms. According to WHO reports from 2014, there has been an alarming increase in the antibiotic resistant bacterial strains in most parts of the world1. Our previous results showed that a nanoantibiotic (NAB) design created in our laboratory2, composed of a cerium oxide core, mesoporous silica shell loaded with capsaicin, and a chitosan coating, are effective against planktonic E. coli. However, most of the pathogenic bacteria form biofilms during infections. That is why the next stage of studying NAB is to determine whether they are effective against biofilms of different species. Moreover, the results of NAB efficiency against planktonic E. coli did not clearly show the contribution of the antibiotic drug component of NAB &#8211; capsaicin. Hence, the first step of the current study is to determine whether and to what degree, mesoporous silica nanoparticles (MSN) &#8211; serving as NAB model in this case - penetrate biofilms as a function of particle shape and surface coating; as well as finding the efficient concentration of capsaicin against E. coli and S. aureus &#160;to optimize the NAB dosing against biofilms. &#160; Aim To check in vitro penetration of MSN on S. aureus biofilm and antibacterial activity of NAB and pure capsaicin on E. coli and S. aureus biofilms. Methods To investigate NAB efficiency on biofilms MBEC-high-throughput assay3 was performed. Equal biofilms formed on peg-lids were incubated with different concentrations of NAB and capsaicin. After different time point biofilms were sonicated and plated on agar plated to perform CFU counting. To determine the efficient concentration of capsaicin, biofilms were formed in 12 well plates and then incubated with different concentrations of capsaicin. To visualize inhibitory effect, plating for CFU counting and Resazurin assay were applied. To evaluate the penetration of particles, labeled and non-labeled particles were added to fully grown St. aureus biofilms, incubated and visualized with confocal microscopy and structured illumination microscopy. &#160; Results <ol> <li>Through two different microscopy techniques penetration of particles into biofilm and their localization next to bacteria cells were observed.</li> <li>In MBEC-high-throughput assay no inhibitory effect of NAB against E. coli biofilms was detected in comparison with untreated bacteria.</li> <li>Resazurin assay and CFU counting method allowed us to determine the most efficient concentration of capsaicin against E. coli and St. aureus biofilms.</li> </ol> &#160; Conclusion <ol> <li>Use of MSN and NAB in particular to deliver active antibacterial agents inside the biofilm is justified.</li> <li>We cannot claim that NAB does not demonstrate any activity against E. coli biofilms, though we can suggest that the peg-lid set up is not sufficient for the NAB design. Further experiments are required.</li> <li>The next step is to test different concentrations of NAB against biofilms with more appropriate methods than MBEC-high-throughput assay. These results will allow us to make conclusions about the benefits of NAB in comparison with pure capsaicin.</li> </ol> &#160; References <ol> <li>Govardhanam, N.P. (2017). Development of nanoantibiotics and evaluation via in vitro and in vivo imaging. University of Turku, Finland.</li> <li>Ventola, C. Lee.&#160;Pharmacy and Therapeutics&#160;40.4: 277, 2015</li> <li>Harrison, J. et al., BMC microbiology 5(1), 53, 2005.</li> </ol>

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Role of F1C Fimbriae, Flagella, and Secreted Bacterial Components in the Inhibitory Effect of Probiotic Escherichia coli Nissle 1917 on Atypical Enteropathogenic E. coli Infection

Infection and Immunity ◽

10.1128/iai.01431-13 ◽

2014 ◽

Vol 82 (5) ◽

pp. 1801-1812 ◽

Cited By ~ 23

Author(s):

Sylvia Kleta ◽

Marcel Nordhoff ◽

Karsten Tedin ◽

Lothar H. Wieler ◽

Rafal Kolenda ◽

...

Keyword(s):

Escherichia Coli ◽

Molecular Mechanisms ◽

Inhibitory Effect ◽

Host Cells ◽

Single Infection ◽

Content Type ◽

E Coli ◽

Initial Adhesion

ABSTRACTEnteropathogenicEscherichia coli(EPEC) is recognized as an important intestinal pathogen that frequently causes acute and persistent diarrhea in humans and animals. The use of probiotic bacteria to prevent diarrhea is gaining increasing interest. The probioticE. colistrain Nissle 1917 (EcN) is known to be effective in the treatment of several gastrointestinal disorders. While bothin vitroandin vivostudies have described strong inhibitory effects of EcN on enteropathogenic bacteria, including pathogenicE. coli, the underlying molecular mechanisms remain largely unknown. In this study, we examined the inhibitory effect of EcN on infections of porcine intestinal epithelial cells with atypical enteropathogenicE. coli(aEPEC) with respect to single infection steps, including adhesion, microcolony formation, and the attaching and effacing phenotype. We show that EcN drastically reduced the infection efficiencies of aEPEC by inhibiting bacterial adhesion and growth of microcolonies, but not the attaching and effacing of adherent bacteria. The inhibitory effect correlated with EcN adhesion capacities and was predominantly mediated by F1C fimbriae, but also by H1 flagella, which served as bridges between EcN cells. Furthermore, EcN seemed to interfere with the initial adhesion of aEPEC to host cells by secretion of inhibitory components. These components do not appear to be specific to EcN, but we propose that the strong adhesion capacities enable EcN to secrete sufficient local concentrations of the inhibitory factors. The results of this study are consistent with a mode of action whereby EcN inhibits secretion of virulence-associated proteins of EPEC, but not their expression.

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Experimental Application of Sage in Rabbit Husbandry

Acta Veterinaria Brno ◽

10.2754/avb200877040581 ◽

2008 ◽

Vol 77 (4) ◽

pp. 581-588 ◽

Cited By ~ 10

Author(s):

R. Szabóová ◽

A. Lauková ◽

Ľ. Chrastinová ◽

M. Simonová ◽

V. Strompfová ◽

...

Keyword(s):

Inhibitory Effect ◽

Negative Influence ◽

Feed Consumption ◽

Control Group ◽

Coagulase Negative Staphylococci ◽

E Coli ◽

Commercial Feed ◽

Experimental Group

Salvia spp. belongs to the Labiatae family and is characterized by antimicrobial and antiinflammatory effect. The aim of this study was to test its in vitro and in vivo inhibitory effect against bacteria as well as to find an alternative possibility to use sage in the rabbit ecosystem examining biochemical, zootechnical and inmunological indicators, compared to the commercial feed mixture Xtract. Using the sage extract in in vitro tests, its inhibitory effect was noted. Under in vivo conditions, in the experimental group with sage (EG1), reduction of Pseudomonas-like sp. (p < 0.01) and E. coli (p < 0.01) was noted after 7 days of sage application compared to the control group CG2 (with Robenidin) as well as after 21 days of sage extract application, when the reduction of coagulase-negative staphylococci (p < 0.01) was detected (in comparison with the experimental group-EG2, Xtract group). In the caecum of rabbits from EG1, higher values of lactic, acetic and butyric acids were noted. The values of propionic acid were not influenced. Biochemical indicators were not influenced; however, the values of GSH Px were lower in EG1 compared to EG2. Higher phagocytic activity (18%) was noted in EG1 than in EG2 (13%) after 21 days of additives application. The reduction of Eimeria sp. oocysts was demonstrated in EG1 (sage group) after 7 days of sage application comparing to CG2 (217 OPG to 566 OPG). The animals in both experimental groups achieved higher feed consumption and weight gain, lower mortality compared to both controls. Neither of the additives had a negative influence on the health status and growth performance of rabbits.

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In Vitro Inactivation of Escherichia coli O157:H7 in Bovine Rumen Fluid by Caprylic Acid

Journal of Food Protection ◽

10.4315/0362-028x-67.5.884 ◽

2004 ◽

Vol 67 (5) ◽

pp. 884-888 ◽

Cited By ~ 26

Author(s):

THIRUNAVUKKARASU ANNAMALAI ◽

MANOJ KUMAR MOHAN NAIR ◽

PATRICK MAREK ◽

PRADEEP VASUDEVAN ◽

DAVID SCHREIBER ◽

...

Keyword(s):

Escherichia Coli ◽

Caprylic Acid ◽

Rumen Fluid ◽

Bacterial Load ◽

Anaerobic Bacteria ◽

Inhibitory Effect ◽

Escherichia Coli O157 ◽

E Coli ◽

Bovine Rumen

The antibacterial effect of caprylic acid (35 and 50 mM) on Escherichia coli O157:H7 and total anaerobic bacteria at 39° C in rumen fluid (pH 5.6 and 6.8) from 12 beef cattle was investigated. The treatments containing caprylic acid at both pHs significantly reduced (P < 0.05) the population of E. coli O157:H7 compared with that in the control samples. At pH 5.6, both levels of caprylic acid killed E. coli O157:H7 rapidly, reducing the pathogen population to undetectable levels at 1 min of incubation (a more than 6.0-log CFU/ml reduction). In buffered rumen fluid at pH 6.8, 50 mM caprylic acid reduced the E. coli O157:H7 population to undetectable levels at 1 min of incubation, whereas 35 mM caprylic acid reduced the pathogen by approximately 3.0 and 5.0 log CFU/ml at 8 and 24 h of incubation, respectively. At both pHs, caprylic acid had a significantly lesser (P < 0.05) and minimal inhibitory effect on the population of total anaerobic bacteria in rumen compared with that on E. coli O157:H7. At 24 h of incubation, caprylic acid (35 and 50 mM) reduced the population of total anaerobic bacteria by approximately 2.0 log CFU/ml at pH 5.6, whereas at pH 6.8, caprylic acid (35 mM) did not have any significant (P > 0.05) inhibitory effect on total bacterial load. Results of this study revealed that caprylic acid was effective in inactivating E. coli O157:H7 in bovine rumen fluid, thereby justifying its potential as a preslaughter dietary supplement for reducing pathogen carriage in cattle.

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In Vitro Antibacterial Activities and Mechanisms of Action of Fatty Acid Monoglycerides Against Four Foodborne Bacteria

Journal of Food Protection ◽

10.4315/0362-028x.jfp-19-259 ◽

2020 ◽

Vol 83 (2) ◽

pp. 331-337

Author(s):

WENYUE WANG ◽

RUI WANG ◽

GUIJU ZHANG ◽

FANGLI CHEN ◽

BAOCAI XU

Keyword(s):

Antibacterial Activity ◽

Antibacterial Activities ◽

Carbon Chain ◽

Antimicrobial Activities ◽

Inhibitory Effect ◽

Gram Positive ◽

Gram Negative ◽

E Coli ◽

Alkaline Conditions

ABSTRACT Naturally occurring monoglyceride esters of fatty acids have been associated with a broad spectrum of antimicrobial activities. We used an automated turbidimetric method to measure the MIC and assess the antimicrobial activity of five monoglycerides (monocaprin, monolaurin, monomyristin, monopalmitin, and monostearin) against pathogenic strains of Staphylococcus aureus, Bacillus subtilis, Pseudomonas aeruginosa, and Escherichia coli. The antibacterial activity of monocaprin was highest because its carbon chain is shorter than those of other monoglycerides. The MICs of monocaprin against S. aureus, B. subtilis, P. aeruginosa, and E. coli were 0.32, 0.32, 2.5, and 2.5 mg/mL, respectively. Monocaprin had antibacterial activity under neutral and alkaline conditions (pH 7.0 to 9.0) but had no inhibitory effect on S. aureus, B. subtilis, and E. coli under weakly acidic conditions (pH 6.0). The antibacterial mechanism of monocaprin against gram-positive strains (S. aureus and B. subtilis) resulted from destruction of the cell membrane. In contrast, the antibacterial activity of monocaprin against gram-negative strains (P. aeruginosa and E. coli) was attributed to damage to lipopolysaccharides in the cell walls. Because of its inhibitory effect on both gram-positive and gram-negative bacteria, monocaprin could be used as an antibacterial additive in the food industry. HIGHLIGHTS

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THE ACTION OF SULFONAMIDES ON THE RESPIRATION OF BACTERIA AND YEAST

The Journal of General Physiology ◽

10.1085/jgp.25.6.805 ◽

1942 ◽

Vol 25 (6) ◽

pp. 805-817 ◽

Cited By ~ 8

Author(s):

M. G. Sevag ◽

M. Shelburne ◽

Stuart Mudd

Keyword(s):

Structural Similarity ◽

Inhibitory Effect ◽

Brewer’S Yeast ◽

Respiratory Enzymes ◽

Staph Aureus ◽

E Coli ◽

Brewer's Yeast ◽

Inhibiting Effect

The inhibiting effects of sulfonamide drugs and their derivatives on the anaerobic decarboxylation of pyruvic acid by Staphylococcus aureus, Escherichia coli, baker's and brewer's yeast, and a carboxylase preparation from brewer's yeast have been investigated. These drugs are: sulfanilamide, sulfapyridine, sulfadiazine, sulfamethyldiazine, sulfathiazole, sulfamethylthiazole, sulfanilamido-5-ethyl-4-thiazolone, 2-aminopyrimidine, 2-aminothiazole, and 2-aminopyridine. The sulfathiazole ring appears to exercise decidedly greater specific inhibiting effect on the carboxylases of Staph. aureus and E. coli. The inhibiting effect on yeast carboxylase is non-differentiable among all the substances tried, except sulfamethyldiazine which is completely ineffective on the carboxylases of the organisms studied. The specific inhibitory effect of sulfathiazole on the carboxylases of Staph. aureus and E. coli in comparison to sulfanilamide, sulfapyridine, and sulfadiazine is in harmony with in vivo and in vitro experimental results of other investigators. The results of the present investigation appear to support the hypothesis (1) that sulfonamides exert their bacteriostatic action through chemical affinity for the carrier proteins of certain respiratory enzymes of the bacterial cell, and that this affinity may in part be related to structural similarity between components of the drugs and the corresponding respiratory coenzymes.

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