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Baron S, editor. Medical Microbiology. Fourth edition. Galveston (TX): college of Texas clinical Branch in ~ Galveston; 1996.


Gross Morphology

Bacteria have characteristic forms (cocci, rods, spirals, etc.) and often occurin properties aggregates (pairs, chains, tetrads, clusters, etc.). Thesetraits room usually usual for a genus and are diagnostically useful.

Cell Structure

Prokaryotes have actually a new closed (nuclear body) quite than one enveloped cell nucleus andlack membrane-bound cytoplasmic organelles. The plasma membrane in prokaryotesperforms countless of the functions lugged out through membranous oribel ineukaryotes. Multiplication is by binary fission.

Surface Structures

Flagella: The flagella the motile bacteria differ in framework fromeukaryotic flagella. A basal body anchored in the plasma membrane and also cell wallgives climb to a cylindrical protein filament. The flagellum move by whirlingabout its lengthy axis. The number and also arrangement the flagella ~ above the cell arediagnostically useful.

Pili (Fimbriae): Pili room slender, hairlike, proteinaceousappendages top top the surface of plenty of (particularly Gram-negative) bacteria. Theyare important in adhesion to hold surfaces.

Capsules: part bacteria type a thick outer capsule ofhigh-molecular-weight, viscous polysaccharide gel; others have an ext amorphousslime layers. Capsules confer resistance to phagocytosis.

Important Chemical materials of surface ar Structures

Cell wall surface Peptidoglycans: Both Gram-positive and Gram-negativebacteria own cell wall peptidoglycans, i m sorry confer the characteristics cellshape and provide the cell with mechanical protection. Peptidoglycans room uniqueto prokaryotes organisms and also consist the a glycan backbone that muramic acid andglucosamine (both N-acetylated), and also peptide chains highly cross-linked withbridges in Gram-positive bacteria (e.g., Staphylococcus aureus)or partly cross-linked in Gram-negative bacteria (e.g., Escherichiacoli). The cross-linking transpeptidase enzymes are few of thetargets for b-lactam antibiotics.

Teichoic Acids: Teichoic acids are polyol phosphate polymers bearinga strong an adverse charge. They space covalently connected to the peptidoglycan insome Gram-positive bacteria. They are strongly antigenic, however are generallyabsent in Gram-negative bacteria.

Lipoteichoic Acids: Lipoteichoic acids as membrane teichoic acidsare polymers of amphiphitic glycophosphates through the lipophilic glycolipid andanchored in the cytoplasmic membrane. They space antigenic, cytotoxic and adhesins(e.g., Streptococcus pyogenes).

Lipopolysaccharides: among the significant components that the outermembrane the Gram-negative bacteria is lipopolysaccharide (endotoxin), a complexmolecule consist of of a lipid A anchor, a polysaccharide core, and chains ofcarbohydrates. Street in the polysaccharide chain confer serologicspecificity.

Wall-Less Forms: Two groups of bacteria there is no of cabinet wallpeptidoglycans room the Mycoplasma species, which own asurface membrane structure, and the L-forms that arise from either Gram-positiveor Gram-negative bacterial cells that have lost their capability to develop thepeptidoglycan structures.

Cytoplasmic Structures

Plasma Membrane: The bacterial plasma membrane is created primarilyof protein and also phospholipid (about 3:1). That performs plenty of functions, includingtransport, biosynthesis, and energy transduction.

Organelles: The bacter cytoplasm is densely packed with 70Sribosomes. Various other granules represent metabolic to make reservation (e.g.,poly-β-hydroxybutyrate, polysaccharide, polymetaphosphate, andmetachromatic granules).

Endospores:Bacillus and also Clostridium species can produceendospores: heat-resistant, dehydrated relaxing cells that space formedintracellularly and also contain a genome and also all necessary metabolic machinery. Theendospore is encased in a facility protective spore coat.


All bacteria, both pathogenic and saprophytic, space unicellular organisms thatreproduce through binary fission. Most bacteria are capable of live independence metabolicexistence and also growth, but species of Chlamydia andRickettsia are obligately intracellular organisms. Bacterialcells room extremely tiny and are most conveniently measure in microns(10-6 m). They variety in size from big cells such asBacillus anthracis (1.0 come 1.3 µm X 3 to 10µm) to very small cells such together Pasteurella tularensis(0.2 X 0.2 to 0.7 µm) Mycoplasmas (atypical pneumoniagroup) are even smaller, measure 0.1 to 0.2 µm in diameter. Bacteriatherefore have a surface-to-volume proportion that is very high: about 100,000.

Bacteria have actually characteristic shapes. The usual microscopic morphologies space cocci(round or ellipsoidal cells, such together Staphylococcus aureus orStreptococcus, respectively); rods, such asBacillus and also Clostridium species; long,filamentous branched cells, such as Actinomyces species; andcomma-shaped and also spiral cells, such together Vibrio cholerae andTreponema pallidum, respectively. The setup of cell isalso usual of various species or groups of bacteria (Fig. 2-1). Part rods or cocci characteristically flourish inchains; some, such as Staphylococcus aureus, form grapelikeclusters of spherical cells; part round cocci form cubic packets. Bacterial cell ofother varieties grow separately. The microscopic appearance is therefore beneficial inclassification and also diagnosis. The higher resolving power of the electron microscopenot only magnifies the typical shape that a bacter cell yet also plainly resolvesits prokaryotic organization (Fig. 2-2).



Figure 2-2

Electron micrograph of a thin ar of Neisseriagonorrhoeae reflecting the organizational features ofprokaryotic cells. Keep in mind the electron-transparent nuclear an ar (n) packed through DNA fibrils,the thick distribution of ribosomal corpuscle in the cytoplasm, (more...)

The Nucleoid

Prokaryotic and also eukaryotic cells were initially distinguished on the communication ofstructure: the prokaryotic nucleoidthe equivalent of the eukaryotic bio nucleusisstructurally much easier than the true eukaryotic nucleus, which has actually a facility mitoticapparatus and surrounding nuclear membrane. As the electron micrograph in Fig. 2-2 shows, the bacter nucleoid, whichcontains the DNA fibrils, lacks a limiting membrane. Under the irradiate microscope, thenucleoid of the bacter cell deserve to be visualized with the help of Feulgen staining,which stains DNA. Gentle lysis deserve to be supplied to isolate the nucleoid of most bacterialcells. The DNA is then viewed to it is in a single, continuous, "giant" one moleculewith a molecular weight of approximately 3 X 109 (see Ch. 5). The unfolded nuclear DNA would certainly beabout 1 mm lengthy (compared with an average size of 1 to 2 µm forbacterial cells). The bacterial nucleoid, then, is a framework containing a singlechromosome. The number of copies that this chromosome in a cell relies on the stageof the cabinet cycle (chromosome replication, cabinet enlargement, chromosome segregation,etc). Back the mechanism of segregation of the 2 sister chromosomes followingreplication is not completely understood, every one of the models proposed call for that thechromosome be permanently attached to the cabinet membrane transparent the variousstages the the cabinet cycle.

Bacterial dyed does not contain straightforward histone proteins, however low-molecular-weightpolyamines and also magnesium ions may meet a role similar to that of eukaryotichistones. Despite the differences between prokaryotic and also eukaryotic DNA,prokaryotic DNA from cells infected v bacteriophage 𝛄, whenvisualized through electron microscopy, has actually a beaded, condensed appearance not unlikethat of eukaryotic bio chromatin.

Surface Appendages

Two types of surface ar appendage deserve to be recognized on specific bacterial species: theflagella, which space organs of locomotion, and also pili (Latin hairs), which are alsoknown together fimbriae (Latin fringes). Flagella take place on both Gram-positive andGram-negative bacteria, and their presence deserve to be beneficial in identification. Forexample, castle are discovered on many species of bacilli but rarely ~ above cocci. In contrast,pili occur nearly exclusively ~ above Gram-negative bacteria and are uncovered on only a fewGram-positive organisms (e.g., Corynebacterium renale).

Some bacteria have both flagella and also pili. The electron micrograph in Fig. 2-3 shows the characteristic wavyappearance the flagella and two varieties of pili on the surface of Escherichiacoli.


Figure 2-3

(A) Electron micrograph of negatively stained E. Colishowing wavy flagella and also numerous short, thinner, and much more rigidhairlike structures, the pili. (B) The lengthy sex pilus have the right to bedistinguished from the shorter common chili by mix E.coli cells v (more...)


Structurally, bacter flagella are lengthy (3 come 12 µm), filamentoussurface appendages about 12 to 30 nm in diameter. The protein subunits the aflagellum room assembled to kind a cylindrical framework with a hole core. Aflagellum consists of three parts: (1) the long filament, i beg your pardon lies outside tothe cabinet surface; (2) the hook framework at the end of the filament; and also (3) thebasal body, to which the hook is anchored and which imparts motion to theflagellum. The basal body traverses the outer wall surface and membrane structures. Itconsists the a rod and one or 2 pairs that discs. The thrust that propels thebacterial cabinet is noted by counterclockwise rotation of the basal body, whichcauses the helically twisted filament come whirl. The motion of the basal bodyis moved by a proton motive pressure rather than by ATP directly. The capacity ofbacteria come swim by way of the propeller-like activity of the flagella providesthem v the mechanical way to execute chemotaxis (movement in solution toattractant and repellent substances in the environment). Solution to chemicalstimuli involves a innovative sensory mechanism of receptors the are situated inthe cell surface and/or periplasm and also that transmit information tomethyl-accepting chemotaxis protein that manage the flagellar motor. Geneticstudies have actually revealed the existence of mutants with altered biochemical pathwaysfor flagellar motility and chemotaxis.

Chemically, flagella are built of a course of proteins dubbed flagellins.The hook and basal-body structures consist of many proteins. Mutationsaffecting any kind of of this gene assets may result in loss or handicap ofmotility. Flagellins are immunogenic and also constitute a team of protein antigenscalled the H antigens, which space characteristic that a given species, strain, orvariant of one organism. The varieties specificity the the flagellins reflectsdifferences in the primary structures of the proteins. Antigenic transforms of theflagella known as the step variation of H1 and H2 wake up in Salmonellatyphimurium (see Ch.21 and Ref. Seifert andSo).

The number and also distribution of flagella top top the bacterial surface arecharacteristic because that a given types and hence are advantageous in identify andclassifying bacteria. Figure 2-4illustrates common arrangements that flagella ~ above or approximately the bacter surface.For example, V. Cholerae has a single flagellum at one pole ofthe cabinet (i.e., it is monotrichous), vice versa, Proteus vulgarisand E. Coli have countless flagella distributed over the whole cellsurface (i.e., they space peritrichous). The flagella the a peritrichous bacteriummust accumulation as a posterior bundle come propel the cabinet in a forwarddirection.


Flagella deserve to be sheared indigenous the cell surface without affecting the viability ofthe cell. The cell then becomes temporarily nonmotile. Gradually it synthesizesnew flagella and also regains motility. The protein synthetic inhibitorchloramphenicol, however, blocks rebirth of flagella.


The state pili and fimbriae space usually used interchangeably to describe thethin, hairlike appendages top top the surface of many Gram-negative bacteria andproteins of pili are referred to as pilins. Pili are much more rigid in appearancethan flagella (Fig. 2-3). In someorganisms, such as Shigella types and E.coli, chili are dispersed profusely over the cell surface, v asmany as 200 every cell. As is quickly recognized in strains the E.coli, pili deserve to come in 2 types: short, abundant common pili, anda little number (one to six) of an extremely long pili well-known as sex pili. Sex pili deserve to bedistinguished by their capability to tie male-specific bacteriophages (the sexpilus acts as a certain receptor because that these bacteriophages) (Fig. 2-3B). The sex pili attach male tofemale bacteria throughout conjugation.

Pili in plenty of enteric bacteria confer adhesive nature on the bacter cells,enabling them to adhere to various epithelial surfaces, to red blood cells(causing hemagglutination), and to surface of yeast and also fungal cells. Theseadhesive properties of piliated cells play vital role in bacterialcolonization the epithelial surfaces and also are as such referred to ascolonization factors. The common pili discovered on E. Coli exhibita sugar specificity analogous to that of phytohemagglutinins and also lectins, inthat adhesion and hemagglutinating capacities that the organism room inhibitedspecifically by mannose. Biology possessing this type of hemagglutination arecalled mannose-sensitive organisms. Other piliated organisms, such together gonococci,are adhesive and hemagglutinating, but are insensitive to the inhibitory effectsof mannose. Substantial antigenic sport in pilins the gonococci space well known(see Ref. Seifert and also So).

Surface Layers

The surface ar layers of the bacterial cell have been determined by various techniques:light microscopy and also staining; electron microscopy of thin-sectioned,freeze-fractured, and also negatively stained cells; and also isolation and biochemicalcharacterization of individual morphologic contents of the cell. The principalsurface layers space capsules and loose slime, the cell wall surface of Gram-positive bacteriaand the complex cell envelope that Gram-negative bacteria, plasma (cytoplasmic)membranes, and also mesosomal membrane vesicles, i m sorry arise indigenous invaginations that theplasma membrane. In bacteria, the cell wall surface forms a rigid structure of uniformthickness roughly the cell and is responsible for the characteristic shape of thecell (rod, coccus, or spiral). Inside the cell wall (or strictly peptidoglycan layer)is the plasma (cytoplasmic) membrane; this is usually carefully apposed to the walllayer. The topographic relationship of the cell wall and envelope layers come theplasma membrane are suggested in the thin section of a Gram-positive organism(Micrococcus lysodeikticus) in figure 2-5A and also in the freeze-fractured cell of a Gram-negative organism(Bacteroides melaninogenicus) in figure 2-5B. The latter shows the common fracture plane seenin many Gram-negative bacteria, which room weak cleavage planes through the outermembrane the the envelope and extensive fracture planes v the bilayer region ofthe basic plasma membrane.

Figure 2-5

(A) Electron micrograph of a thin ar of the Gram-positive M.lysodeikticus reflecting the special peptidoglycan cabinet wall(cw), basic cytoplasmic (plasma) membrane (cm), mesosome (m), andnucleus (n). (B) Freeze-fractured Bacteriodes cellshowing (more...)

Capsules and loose Slime

Some bacteria kind capsules, i m sorry constitute the outermost layer of thebacterial cell and also surround it v a reasonably thick layer of viscous gel.Capsules may be as much as 10 µm thick. Some organisms absence a well-definedcapsule however have loose, amorphous slime layers exterior to the cell wall or cellenvelope. The α hemolytic Streptococcus mutans, theprimary organism found in dentist plaque is maybe to synthesis a largeextracellular mucoid glucans indigenous sucrose. No all bacterial species producecapsules; however, the capsules the encapsulated pathogens are regularly importantdeterminants of virulence. Encapsulated species are found amongst bothGram-positive and also Gram-negative bacteria. In both groups, most capsules arecomposed the highmolecular-weight viscous polysaccharides the are retained as athick gel external the cell wall or envelope. The capsule that Bacillusanthracis (the causal agent of anthrax) is inexplicable in the it iscomposed the a γ-glutamyl polypeptide. Table 2-1 presents the various capsular building material formedby a choice of Gram-positive and also Gram-negative bacteria. A plasma membranestage is affiliated in the biosynthesis and also assembly the the capsular substances,which space extruded or secreted with the outer wall or envelope structures.Mutational ns of enzymes affiliated in the biosynthesis the the capsularpolysaccharides can result in the smooth-to-rough variation watched in thepneumococci.

The capsule is not vital for viability. Viability is not affected whencapsular polysaccharides are eliminated enzymatically indigenous the cell surface. Theexact attributes of capsules are not completely understood, but they perform conferresistance come phagocytosis and also hence administer the bacter cell through protectionagainst hold defenses come invasion.

Cell wall and Gram-Negative cabinet Envelope

The Gram stain generally differentiates bacteria right into Gram-positive andGram-negative groups; a few organisms are repeatedly Gram-variable.Gram-positive and also Gram-negative organisms differ drastically in the organizationof the structures exterior the plasma membrane but below the capsule (Fig. 2-6): in Gram-negative biology thesestructures constitute the cell envelope, whereas in Gram-positive biology theyare referred to as a cabinet wall.

Figure 2-6

Comparison the the special cell wall of Gram-positive bacteria withthe comparatively slim cell wall of Gram-negative bacteria. Note the complexity of the Gram-negative cabinet envelope (outermembrane, that hydrophobic lipoprotein anchor; periplasmicspace). (more...)

Most Gram-positive bacteria have a fairly thick (about 20 to 80 nm),continuous cell wall surface (often called the sacculus), which is composed greatly ofpeptidoglycan (also well-known as mucopeptide or murein). In thick cell walls, othercell wall surface polymers (such as the teichoic acids, polysaccharides, andpeptidoglycolipids) are covalently attached come the peptidoglycan. In contrast,the peptidoglycan layer in Gram-negative bacteria is thin (about 5 come 10 nmthick); in E. Coli, the peptidoglycan is more than likely only amonolayer thick. External the peptidoglycan class in the Gram-negative envelopeis an external membrane structure (about 7.5 to 10 nm thick). In most Gram-negativebacteria, this membrane structure is anchored noncovalently come lipoproteinmolecules (Braun"s lipoprotein), which, in turn, are covalently connected to thepeptidoglycan. The lipopolysaccharides that the Gram-negative cell envelope formpart of the outer leaflet the the external membrane structure.

The organization and overall size of the external membrane that theGram-negative cell envelope are similar to those of the plasma membrane (about7.5 nm thick). Moreover, in Gram-negative bacteria such as E.coli, the outer and inner membranes adhere come each various other at severalhundred web page (Bayer patches); this sites have the right to break increase the continuity of thepeptidoglycan layer. Table 2-2summarizes the significant classes of chemistry constituents in the walls and envelopesof Gram-positive and Gram-negative bacteria.

The basic differences in surface structures of Gram-positive and also Gram-negativebacteria explain the results of Gram staining. Both Gram-positive andGram-negative bacteria take it up the same amounts of decision violet (CV) andiodine (I). The CV-I complex, however, is trapped within the Gram-positive cellby the dehydration and reduced porosity of the thick cell wall as a result ofthe differential washing action with 95 percent ethanol or other solvent mixture.In contrast, the slim peptidoglycan layer and probable discontinuities at themembrane adhesion sites carry out not impede solvent extraction of the CV-I complexfrom the Gram-negative cell. The above mechanism that the Gram stain based upon thestructural differences between the two teams has been evidenced bysophisticated methods of electron microscopy (see Ref. Bereridge and Daries).The sequence of actions in the Gram stain differentiation is illustrateddiagrammatically in figure 2-7. Moreover,mechanical disruption that the cell wall surface of Gram-positive organisms or itsenzymatic removal through lysozyme outcomes in complete extraction of the CV-Icomplex and conversion to a Gram-negative reaction. Therefore, autolyticwall-degrading enzymes that reason cell wall breakage might account forGram-negative or variable reactions in cultures of Gram-positive organisms (suchas Staphylococcus aureus, Clostridiumperfringens, Corynebacterium diphtheriae, and also someBacillus spp.).

Figure 2-7

General sequence of procedures in the Gram stain procedure and theresultant staining the Gram-positive and Gram-negativebacteria.


Unique attributes of virtually all prokaryotes cells (except for Halobacteriumhalobium and also mycoplasmas) room cell wall peptidoglycan and thespecific enzymes connected in the biosynthesis. This enzymes room target sitesfor inhibition the peptidoglycan synthesis by specific antibiotics. The primarychemical structures of peptidoglycans of both Gram-positive and also Gram-negativebacteria have actually been established; they consist of a glycan backbone the repeatinggroups the β1, 4-linked disaccharides ofβ1,4-N-acetylmuramyl-N-acetylglucosamine. Tetrapeptides ofL-alanine-D-isoglutamic acid-L-lysine (or diaminopimelic acid)-n-alanine arelinked v the carboxyl team by amide link of muramic acid residues ofthe glycan chains; the D-alanine residues are directly cross-linked to the𝛆-amino team of lysine or diaminopimelic acid on a neighboringtetrapeptide, or they are linked by a peptide bridge. In S.aureus peptidoglycan, a glycine pentapeptide bridge links the twoadjacent peptide structures. The degree of straight or peptide-bridgecross-linking different from one peptidoglycan to another. The staphylococcalpeptidoglycan is highly cross-linked, whereas the of E. Coliis much much less so, and also has a much more open peptidoglycan mesh. The diamino acidproviding the 𝛆-amino team for cross-linking is lysine ordiaminopimelic acid, the latter being uniformly current in Gram-negativepeptidoglycans. The structure of the peptidoglycan is portrayed in figure 2-8. A peptidoglycan with a chemicalstructure substantially different from that of all eubacteria has beendiscovered in certain archaebacteria. Rather of muramic acid, thispeptidoglycan consists of talosaminuronic acid and also lacks the D-amino acids found inthe eubacterial peptidoglycans. Interestingly, biology containing this wallpolymer (referred to as pseudomurein) room insensitive to penicillin, aninhibitor the the transpeptidases affiliated in peptidoglycan biosynthesis ineubacteria.

Figure 2-8

Diagrammatic representation of peptidoglycan frameworks withadjacent glycan strands cross-linked directly from thecarboxyterminal D-alanine to the 𝛆-amino team of anadjacent tetrapeptide or v a peptide cross bridge,N-acetylmuramic acid, (more...)

The ß-1,4 glycosidic bond in between N-acetylmuramic acid andN-acetylglucosamine is particularly cleaved through the bacteriolytic enzymelysozyme. Widely spread in nature, this enzyme is current in human tissuesand secretions and also can cause complete digestion of the peptidoglycan walls ofsensitive organisms. Once lysozyme is allowed to digest the cell wall surface ofGram-positive bacteria rely in one osmotic stabilizer (such as sucrose),protoplasts are formed. This protoplasts are able come survive and continue togrow on suitable media in the wall-less state. Gram-negative bacteria treatedsimilarly produce spheroplasts, i m sorry retain lot of the outer membranestructure. The dependence of bacterial shape on the peptidoglycan is displayed bythe revolution of rod-shaped bacteria to spherical protoplasts(spheroplasts) after enzymatic malfunction of the peptidoglycan. The mechanicalprotection afforded by the wall peptidoglycan layer is apparent in the osmoticfragility that both protoplasts and spheroplasts. There are two groups of bacteriathat lack the security cell wall peptidoglycan structure, the Mycoplasmaspecies, among which causes atypical pneumonia and some genitourinary tractinfections and also the L-forms, which originate indigenous Gram-positive or Gram-negativebacteria and are for this reason designated since of their discovery and description at theLister Institute, London. The mycoplasmas and also L-forms room all Gram-negative andinsensitive come penicillin and are bounded through a surface ar membrane structure.L-forms occurring "spontaneously" in societies or diverted from epidemic arestructurally concerned protoplasts and also spheroplasts; all 3 forms(protoplasts, spheroplasts, and L-forms) revert infrequently and only underspecial conditions.

Teichoic Acids

Wall teichoic acids are discovered only in details Gram-positive bacteria (such asstaphylococci, streptococci, lactobacilli, and also Bacillus spp.);so far, they have not been uncovered in gram- an unfavorable organisms. Teichoic acids arepolyol phosphate polymers, with either ribitol or glycerol linked byphosphodiester bonds; their frameworks are shown in figure 2-9. Substituent teams on the polyol chains caninclude D-alanine (ester linked), N-acetylglucosamine, N-acetylgalactosamine,and glucose; the substituent is characteristic for the teichoic mountain from aparticular bacterial types and have the right to act together a specific antigenic determinant.Teichoic acids space covalently attached to the peptidoglycan. This highlynegatively charged polymers the the bacterial wall can serve as acation-sequestering mechanism.

Figure 2-9

Structures of cell wall surface teichoic acids. (A) Ribitol teichoic acid with repeating units of 1,5-phosphodiesterlinkages the D-ribitol and D-alanyl ester on position 2 and also glycosylsubstituents (R) on position 4. The glycosyl teams may abeN-acetylglucosaminyl (more...)

Accessory wall surface Polymers

In enhancement to the major cell wall polymers, the wall surfaces of certainGram-positive bacteria own polysaccharide molecules linked to thepeptidoglycan. Because that example, the C polysaccharide of streptococci confers groupspecificity. Acidic polysaccharides attached to the peptidoglycan space calledteichuronic acids. Mycobacteria have peptidoglycolipids, glycolipids, and waxesassociated with the cell wall.


A characteristic attribute of Gram-negative bacteria is possession of miscellaneous typesof complicated macromolecular lipopolysaccharide (LPS). Therefore far, only oneGram-positive organism, Listeria monocytogenes, has been foundto contain an really LPS. The LPS of this bacterium and those that allGram-negative species are also called endotoxins, thereby distinguishing thesecell-bound, heat-stable toxin from heat-labile, protein exotoxins secreted intoculture media. Endotoxins possess range of an effective biologic tasks andplay crucial role in the pathogenesis of plenty of Gram-negative bacterialinfections. In enhancement to causing endotoxic shock, LPS is pyrogenic, canactivate macrophages and also complement, is mitogenic for B lymphocytes, inducesinterferon production, causes tissue necrosis and also tumor regression, and also hasadjuvant properties. The endotoxic nature of LPS reside mainly in the lipidA components. Usually, the LPS molecules have three regions: the lipid Astructure compelled for insertion in the external leaflet the the external membranebilayer; a covalently attached core composed of 2-keto-3deoxyoctonic mountain (KDO),heptose, ethanolamine, N-acetylglucosamine, glucose, and galactose; andpolysaccharide chains connected to the core. The polysaccharide chain constitutethe O-antigens that the Gram-negative bacteria, and the separation, personal, instance monosaccharideconstituents confer serologic specificity on these components. Figure 2-10 depicts the structure of LPS.Although it has been known that lipid A is created of β1,6-linkedD-glucosamine disaccharide substituted with phosphomonester groups at positions4" and 1, uncertainties have existed around the attachment location of the sixfatty mountain acyl and KDO teams on the disaccharide. The demonstrate of thestructure of lipid A of LPS that a heptoseless mutant that Salmonellatyphimurium has created that amide-linked hydroxymyristoyl andlauroxymyristoyl groups are attached to the nitrogen that the 2- and also 2"-carbons,respectively, and that hydroxymyristoyl and myristoxymyristoyl groups areattached to the oxygen that the 3- and also 3"-carbons that the disaccharide,respectively. Therefore, just position 6" is left for attachment of KDOunits.

LPS and also phospholipids aid confer asymmetry come the outer membrane the theGram-negative bacteria, with the hydrophilic polysaccharide chains outermost.Each LPS is organized in the outer membrane by relatively weak cohesive forces (ionicand hydrophobic interactions) and also can be dissociated native the cell surface ar withsurface-active agents.

As in peptidoglycan biosynthesis, LPS molecules are assembled in ~ the plasma orinner membrane. These newly created molecules are at first inserted into theouter-inner membrane adhesion sites.

Outer Membrane the Gram-Negative Bacteria

In slim sections, the external membranes of Gram-negative bacteria appear broadlysimilar to the plasma or within membranes; however, they differ from the innermembranes and walls that Gram-positive bacteria in countless respects. The lipid Aof LPS is placed with phospholipids to develop the external leaflet the the bilayerstructure; the lipid part of the lipoprotein and also phospholipid form the innerleaflet the the external membrane double layer of many Gram-negative bacteria (Fig. 2-6).

In addition to these components, the external membrane own several major outermembrane proteins; the most abundant is referred to as porin. The assembled subunits ofporin kind a channel that borders the passage of hydrophilic molecules throughout theouter membrane barrier to those having actually molecular weights that room usually lessthan 600 to 700. Evidence likewise suggests that hydrophobic pathways exist acrossthe external membrane and also are partly responsible because that the differential penetrationand performance of specific b-lactam antibiotics (ampicillin, cephalosporins)that are energetic against miscellaneous Gram-negative bacteria. Return the outermembranes act together a permeability barrier or molecule sieve, they do not appearto possess energy-transducing systems to drive active transport. Number of outermembrane proteins, however, are involved in the details uptake that metabolites(maltose, vitamin B12, nucleosides) and also iron native the medium. Thus, outermembranes the the Gram-negative bacteria carry out a selective barrier to externalmolecules and thereby avoid the loss of metabolite-binding proteins andhydrolytic enzymes (nucleases, alkaline phosphatase) discovered in the periplasmicspace. The periplasmic room is the region between the outer surface the theinner (plasma) membrane and the inner surface of the external membrane (Figure 2-6). Thus, Gram-negative bacteriahave a to move compartment that has no tantamount in Gram-positive organisms.In enhancement to the hydrolytic enzymes, the periplasmic space holds bindingproteins (proteins that particularly bind sugars, amino acids, and also inorganicions) involved in membrane transport and also chemotactic receptor activities.Moreover, plasmid-encoded b-lactamases and also aminoglycoside-modifying enzymes(phosphorylation or adenylation) in the periplasmic room produce antibioticresistance by degrading or editing and enhancing an antibiotic in transit come its targetsites on the membrane (penicillin-binding proteins) or on the ribosomes(aminoglycosides). This periplasmic proteins deserve to be exit by subjecting thecells come osmotic shock and also after treatment v the chelating agentethylenediaminetetraacetic acid.

Plasma (Cytoplasmic) Membranes

Bacterial plasma membranes, the sensible equivalents of eukaryotic plasmamembranes, are referred to variously as cytoplasmic, protoplast, or (inGram-negative organisms) inside membranes. Comparable in in its entirety dimensions andappearance in slim sections come biomembranes from eukaryotic bio cells, lock arecomposed mostly of proteins and also lipids (principally phospholipids).Protein-to-lipid ratios of bacterial plasma membranes are about 3: 1,close come those for mitochondrial membranes. Unlike eukaryotic bio cell membranes,the bacterial membrane (except because that Mycoplasma types andcertain methylotrophic bacteria) has actually no sterols, and bacteria lack the enzymesrequired because that sterol biosynthesis.

Although their composition is comparable to that of inner membranes of Gram-negativespecies, cytoplasmic membranes from Gram-positive bacteria own a course ofmacromolecules not current in the Gram-negative membranes. Plenty of Gram-positivebacterial membranes contain membrane-bound lipoteichoic acid, and specieslacking this component (such together Micrococcus andSarcina spp.) contain an analogous membrane-boundsuccinylated lipomannan. Lipoteichoic acids room structurally comparable to the cellwall glycerol teichoic acids in that they have basal polyglycerol phosphodiester1-3 attached chains (Fig. 2-9). Thesechains terminate v the phosphomonoester end of the polymer, i m sorry is linkedcovalently to one of two people a glycolipid or a phosphatidyl glycolipid moiety. Thus, ahydrophobic tail is provided for anchoring in the membrane lipid layers (Fig. 2-6A). Together in the cell wall surface glycerolteichoic acid, the lipoteichoic acids deserve to have glycosidic and also D-alanyl estersubstituents top top the C-2 position of the glycerol.

Both membrane-bound lipoteichoic acid and membrane-bound succinylated lipomannancan it is in detected together antigens on the cabinet surface, and the glycerol-phosphate andsuccinylated mannan chains show up to prolong through the cell wall structure(Fig. 2-6). This course of polymer hasnot however been uncovered in the cytoplasmic membranes of Gram-negative organisms. Inboth instances, the lipoteichoic acids and the lipomannans room negativelycharged components and also can sequester positively fee substances. Lock havebeen implicated in adhesion to hold cells, but their features remain come beelucidated.

Multiple functions are performed by the plasma membrane of both Gram-positiveand Gram-negative bacteria. Plasma membranes room the website of active transport,respiratory chain components, energy-transducing systems, theH+-ATPase that the proton pump (see thing 4), and membrane step in the biosynthesis ofphospholipids, peptidoglycan, LPS, and also capsular polysaccharides. In essence, thebacterial cytoplasmic membrane is a multifunction framework that combines themitochondrial transport and also biosynthetic features that space usuallycompartmentalized in discrete membranous oribelles in eukaryotic cells. Theplasma membrane is also the anchoring website for DNA and provides the cell with amechanism (as however unknown) because that separation that sister chromosomes.


Thin sections of Gram-positive bacteria expose the visibility of vesicular ortubular-vesicular membrane structures referred to as mesosomes, which room apparentlyformed by one invagination of the plasma membrane. These frameworks are lot moreprominent in Gram-positive 보다 in Gram-negative organisms. In ~ one time, themesosomal motor were assumed to be indistinguishable to bacterial mitochondria;however, numerous other membrane features have additionally been attributed come themesosomes. In ~ present, there is no satisfactory evidence to imply that theyhave a distinctive biochemical or physiologic function. Indeed, electron-microscopicstudies have said that the mesosomes, as commonly seen in thin sections, mayarise from membrane perturbation and fixation artifacts. No basic agreementexists about this theory, however, and some evidence indicates that mesosomesmay be related to occasions in the cell department cycle.

Other Intracellular Components

In enhancement to the nucleoid and cytoplasm (cytosol), the intracellularcompartment that the bacterial cell is densely packed with ribosomes of the 70Stype (Fig. 2-2). This ribonucleoproteinparticles, which have a diameter that 18 nm, are not i ordered it on a membranousrough absorbent reticulum together they space in eukaryotic bio cells. Other granularinclusions randomly spread in the cytoplasm of various species includemetabolic reserve corpuscle such as poly-β-hydroxybutyrate (PHB),polysaccharide and also glycogen-like granules, and polymetaphosphate ormetachromatic granules.

Endospores are very heat-resistant, dehydrated relaxing cells formedintracellularly in members of the genera Bacillus andClostridium. Sporulation, the process of formingendospores, is an unusual building of certain bacteria. The collection ofbiochemical and also morphologic alters that occur throughout sporulation represent truedifferentiation within the bike of the bacterial cell. The process, whichusually starts in the stationary phase of the vegetative cabinet cycle, isinitiated by depletion of nutrient (usually readily utilizable resources ofcarbon or nitrogen, or both). The cell then undergoes a highly complex,well-defined sequence of morphologic and biochemical occasions that ultimately leadto the development of mature endospores. As numerous as seven distinct stages havebeen well-known by morphologic and biochemical research studies of sporulatingBacillus species: stage 0, vegetative cells v twochromosomes at the end of exponential growth; stage I, development of axialchromatin filament and excretion the exoenzymes, including proteases; phase II,forespore septum formation and also segregation of atom material right into twocompartments; stage III, spore protoplast formation and also elevation oftricarboxylic acid and also glyoxylate bike enzyme levels; phase IV, cortexformation and also refractile appearance of spore; phase V, spore coat proteinformation; stage VI, spore maturation, modification of cortical peptidoglycan,uptake the dipicolinic acid (a distinct endospore product) and calcium, anddevelopment of resistance come heat and organic solvents; and stage VII, finalmaturation and liberation the endospores from mommy cells (in part species).

When recently formed, endospores appear as round, extremely refractile cell within thevegetative cell wall, or sporangium. Part strains produce autolysins that digestthe walls and also liberate free endospores. The spore protoplast, or core, containsa finish nucleus, ribosomes, and also energy generating components that areenclosed within a amendment cytoplasmic membrane. The peptidoglycan spore wallsurrounds the spore membrane; on germination, this wall becomes the vegetativecell wall. Neighboring the spore wall surface is a special cortex that contains an unusualtype the peptidoglycan, i m sorry is quickly released top top germination. A spore coat ofkeratinlike protein encases the spore contained within a membrane (theexosporium). During maturation, the spore protoplast dehydrates and also the sporebecomes refractile and resistant to heat, radiation, pressure, desiccation, andchemicals; these properties correlate v the cortical peptidoglycan and thepresence of huge amounts that calcium dipicolinate.

Recent evidence indicated that the spores that Bacillus spharicuswere revitalized which had actually been kept in amber for much more than 25 million years.Their cases need to it is in reevaluated. Figure2-11 illustrates the principal structural functions of a typicalendospore (Bacillus megaterium) on initiation the thegermination process. The thin section of the spore reflects the ruptured, thickspore coat and the cortex neighboring the spore protoplast through the germinalcell wall that i do not care the vegetative wall on outgrowth.

Figure 2-11

Electron micrograph that a thin section of a Bacillusmegaterium spore reflecting the thick spore coat (SC),germinal groove (G) in the spore coat, outer cortex class (OCL) andcortex (Cx) germinal cell wall surface layer (GCW), basic sporeprotoplast membrane (more...)


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light A, Tipper DJ: The outer membrane ofgram-negative bacteria. P. 427. In Sokatch JR, Ornston LN (eds): The bacteria.Vol. 7. Scholastic Press, mountain Diego, 1979 .