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Muramic acid subunit of the peptidoglycan of Escherichia. coli. This is the type of peptidoglycan found in most Gram-negative bacteria. The glycan backbone is a repeat polymer of two amino sugars, N-acetylglucosamine (GlNAc) and N-acetylmuramic acid (MurNAc). Attached to the MurNAc is a tetrapeptide consisting of L-ala-D-glu-DAP-D-ala. In the polymeric form of the molecule, nearby tetrapeptide side chains may be linked to one another by an interpeptide bond between DAP on one chain and D-ala on the other. >  27`Z`Z`Z  + +5  : 0 7H @ 0޽h ? ̙33 H(  HX H c 0A gncwP:  H 08p p( !Figure 9a. Schematic illustration of the outer membrane, cell wall and plasma membrane of a Gram-negative bacterium. Note the structure and arrangement of molecules that constitute the outer membrane. " 2`ZH H 0޽h ? ̙33     P" (  Px P  0̔8S"   8 8Y|(LPS,lipopolysaccharide)P& && $.l P 68   `(A:NNy|x 1uN-YNp S|S|0xxNYyx~b [/f~̃Qk }v;NbR0T1&#"""".# P 6`8  p8h_Y|1u2-n-3-1'l|xKDO)0L-ul-D-u2^|0JSsN|S S|ُ7hN~|{|~b0[NǏKDOkWޏc(W{|A N SNǏa|kWNO-Ovޏ0:X&S""d  P 6@8 J LO-Oyr_'`Y| /f1uY|~bv͑ YUSMO ~g YBg MOÑSOvYhb SỹSObS01uNTyipQl4'`̃Y|vy{|0cRz^TzzgW N T bipQl4'`̃vO-bS g N Tvyr_'`0Da &H" ""$R r P 0ZH ,$D 0 P 08P P v&LPSRP[N,/f1u NR:Sk ~b(2  P 0Ե80 P ( 2\  P 08330  xG-̃vY|bRwQ gk'` 1uN[~̃hb'}[~T S g(W̃SOˆeMbʑ>eQeg Eey:NQk }endotoxin) 02=8n ,.. H P 0޽h ? ̙33*  Xj(  XV X c .A lps  X 08 P tFigure 9b. Structure of LPS " 2`ZH X 0޽h ? ̙33  =5`(  ` ` 680Zg # Figure . Structure of LPS2$ `Z V ` c .A flpsI `  <p8f3 S" P  8 pLPS,lipopolysaccharide& f3 ` s A  ?http://www.liunet.edu/cwis/bklyn/acadres/facdev/FacultyProjects/WebClass/micro-web/images/gm-ve.gifYH ` 0޽h ? ̙33p   h(  h h  68S" s  8 @LPSB\v;NR h 08 W ,`$gbgNipQl4'`~̃uir( Qk }vir(W@x a$w~̃bOb\O(u [SN;bkñv0bu }TgeI{OeQ̃SO _NSN;bkhT(zz-NvvYo b$\O:N͑vbSVP[Q[NipQl4'`̃bSvY7h'` c$/fYlṼSO8TDvSSO0t 2t tH h 0޽h ? ̙33N    p (  p p B8` @Z  E   p B8  E   r  p# #"@ p;  p <8? P  >+Tϑ_NO5~20 e +Tϑؚ~20 +Tϑؚ    @` p <8?P  H+Tϑ_ؚ30~95 +Tϑؚ<50 N,e<2 e%% % @` p <d8?P  & Z| xXx {|( ˆ}v(  @` p <8? 0 P  jipQl4'`~̃  @`  p <8c8?0 P  jipQl3'`~̃  @`  p <0e8?0  6 `S~ހXr^͑v%  @`  p <s8?P  t bR    @``B  p 01 ?ZB  p s *1 ?P P `B p 01 ?`B p 01 ?ZB p s *1 ?`B p 01 ?ZB p s *1 ?0 0 ZB p s *1 ? 0 0 p  0Tw8S" @   8 2Y4ZipQl3'`~̃NipQl4'`~̃~ހXbRk2GG H p 0޽h ? ̙33A0  //__x/(  xA.L ?K x# -N <H x <HmN 4 x 4 x <84 9 Property Z x s *4qN 4 x 4 x <p84 = Gram-positiveZ  x s *4qN <4  x <4  x <8<4 = Gram-negativeZ  x s *<4vN 4  x 4 x <84 BThickness of wallZ x s *4tN 4 x 4 x <#84 @thick (20-80 nm)Z x s *4pN 4< x 4< x <x84< < thin (10 nm) Z x s *4<uN  x  x <x8 ANumber of layersZ x s *xN  x  x < 48 D1 Z x s *wN < x < x <@78< C2 Z x s *<N  x   x <t:8 {Peptidoglycan (murein) content $  Z !x s *zN  "x  #x <=8 F>50% Z $x s *|N < %x < &x <0A8< H10-20% Z 'x s *<N  (x  )x <dD8 aTeichoic acids in wallZ *x s *kN  +x  ,x <G8 7presentZ -x s *jN < .x < /x < K8< 6absentZ 0x s *<N x 1x x 2x <x NLipid and lipoprotein contentZ 3x s *xzN x 4x x 5x <x F0-3% Z 6x s *xyN <x 7x <x 8x <p<x E58% Z 9x s *<xtN x :x x ;x < x @Protein contentZ x < x C0 Z ?x s *xxN x< @x x< Ax <xx< D9% Z Bx s *x<N ` Cx ` Dx <4` eLipopolysaccharide content Z Ex s *`xN ` Fx ` Gx <,` D0 Z Hx s *`yN <` Ix <` Jx </<` E13% Z Kx s *<`N ` Lx ` Mx <3` LSensitivity to Penicillin GZ Nx s *`gN ` Ox ` Px <9` 3yesZ Qx s *`jN `< Rx `< Sx <,=`< 6no (1)Z Tx s *`<N H Ux H Vx <CH iSensitivity to lysozymeZ Wx s *HgN H Xx H Yx <HH 3yesZ Zx s *HjN <H [x <H \x <K<H 6no (2)Z ]x s *<HZ ^x s *h+?K _x 0HGP0J FTable 5. Correlation of Grams stain with other properties of Bacteria.$G F`ZFH x 0޽h ? ̙33 LD(     0 D9S" p  9 ^Y5Z~ހXvR: &$   0xP90% @10Q[NipQlgrv'`( 20Q[~̃vW,gb_` 30Q[~ހvbSOb~ހMQSnSSSv4xOW 40Q[[ñvvOea'` 50Q[N[R }vb'` 60:NkЏRcO/ep 70Q[~ހvbS'` 80Q[~̃vk'`u'` R{ 22 -F1*!0 !0;C!0JQH  0޽h ? ̙33T 0    (  ^ S      c $p! @   2Z|+Tϑؚ `SX͑v30~70%  N T̃y-NZ|()~R N T,wQ͑R{|aIN %ipQl3'`~̃Z|peptidoglycan v~g {^opGr015.016.017.018 NStaphylococcus aureus:NNh0Z|B\S^:N20~80nm 1u~40B\QrRP[~b0QrvZ|'YRP[/f1u'Yϑ\RP[USSOZT bv0kNZ|USSO+T g N*N~bR a) S|USMO N-YNpa|NN-YNpހXxRP[ǏB-1 4-|.ޏc b b) w>\ 1uV*N(lWxޏwegvwޏc(WN-YNpހXxRP[ N0ُV*N(lWx/fL-N(lx-D-7(lx-L-V(lx-D-N(lx c) eh S. Aureusveh:Nu(lxN0ehv(lWzNMRNZ|-NN*N>\v,{V(lWx  D-N(lxvWvޏc [vWzRNvvZ|-NN*N>\v,{ N(lWx  x'`(lWxL-V(lxv(lWvޏc N OMRT$N*NZ|NTweg0 ñvA. Fleming 1922t^Ss X[(WNuSn0NvlmT;m0R~̃TlṼSOQ gHe0W4l~̃Z| \O(uNZ|g NvN-YNpހXxvC1NN-YNpa|C4KNvB-1 4-|.0 Gram positive cells have only two layers, the cytoplasmic membrane and a THICK LAYER of peptidoglycan as the outermost component.-"  0   $+ 2I .0; 5  $+2  !0,H  0޽h ? ̙330 0B(  ^ S    9 c $? @  9 8&{^opGr019.020 :NipQl3'`~̃yr g0 g$Ny{|W a) XxXx NZ|RP[KNSuqQN~T SN(uzxbzxcS +TϑS~ހX͑ϑv50%~ހr^͑v10% +TϑY\NW{QWbR[RvsQ b) xXxsSxXx gulxxRP[N~ހ NvxۏLqQN~TTb_bv vQ+TϑNW{QagNsQ| N'Y SN(u45%pZ4lcS _NSN(up4lN1vQr^~̃-NcS0 xXxv~g;N gulxXxT8h|xXxI{iry{|W ulxXx X[(WNLactobacillus caseir^jsNFg̃ I{~̃-N 8h|xXxX[(WNStaphylococcus aureusTBacillusb[Fg̃^\ I{~̃-N0N,Nỹ+T gNyxX0 F%3$Gw H  0޽h ? ̙33<0 @$(  $^ $S     $c $[ @   \xXxHQ/f(W~ހX-NSsv яt^eg؏Ss(W~ހ(_NX[(W gxXx @bNT Sb:NxXx /H $ 0޽h ? ̙330 P,((  ,^ ,S     ,c $H` @   Another structure in the G+ cell wall is teichoic acid. It is a polymer of glycerol or ribitol joined by phosphate groups. Amino acids, such as D-alanine are attached. Teichoic acid is covalently linked to muramic acid and links various layers of the peptidoglycan mesh together. )4& !0@GH , 0޽h ? ̙3360 `4(  4^ 4S     4c $n @   |:xXxv;NutR a) &^5uw SNsX-NMg++I{3yP[~T cُؚNyP[vSm^ NO~ހ NNNTbv~cؚ;m'`v b) OipQl3'`ũYAet̃ NvQ[;Nv|ޏ;N:NxXx  c) KNipQl3'`̃Nyr_vhbbS d) cOgNlṼSONyr_v8TDSSO0    &  ' H 4 0޽h ? ̙33:0 p<(  <^ <S     <c $ @    {^opGr023 NE. Coli:NNh Z|+Tϑ N~ހXv10% N,1u1~2B\QrRP[gb (W~ހX NvS^N:N~3nm0~gUSSONipQl3'`~̃W,gv T ]+RN(WN a) >\v,{ N*N(lWx:NQmeN(lW^Nxm-DAP  b) l gyrkveh vQMRT$NagZ|KNvT|N1u2u>\,{V(lWx  D-N(lxvWNYN>\,{ N(lWx  m-N(lW^Nxv(lWvcޏc b0 {^opGr023.022 :yipQl3'`~̃N4'`~̃Z|Qv~gT[^v]+R0 YXˆ}v :Nyr_'`}SO,S\Vit B12ُ{|'YRP[eQ~ހQ0 W(ˆ}v  WL](WYXB\-N,YT[ˆ}v, NZSOgbvu4lT[S/zYXB\,O\N800-900DvRP[SǏ,wRP[[{\O(u0 ˆ}v NZ|B\qQN~T,v^Wn(WYXB\,OYXB\brVvNQXB\ޏc0 ipQl4'`~̃vYXB\-N gYvL]ˆ}v( ;N g a) W(ˆ}vmattix protein YE. Coli-NvT[ˆ}vporin 1\/fNyxvzYvW(ˆ}v0T[ˆ}v/fNy NZSO~g kN*NNUSMOvRP[ϑ:N36,00001u NZSO~ggbvEQ4lT[S*jYXB\ SǏRP[ϑ\N800~900vN4l'`%{Qir( OY|{|$\vQ/fS| 0(lWx0N0 N0R }Te:gyP[I{0[OYXB\wQ gRP[[{vR b) YXˆ}vouter membrane protein /fNyyr_'`vЏˆ}vbSSO Sb'YvRP[Y~u }B120[|08hxMirNSyP[TBR  r }ferrichrome bT }enterochelin I{eQ~ހQ c) ˆ}vlipoprotein y{|_Y ;N/fRP[ϑ:N7200vˆ}v vQ\O(u/fO~ހXvYXB\brV0Wޏc(W1uZ|~bvQXB\ N0 In #GRAM NEGATIVE cells the cell wall consists of three components; the cytoplasmic membrane, a thin layer of a rigid sugar MOLECULE NET immediately exterior to the cytoplasmic membrane that covers the entire cytoplasmic membrane and an outermost lipid bilayer called the OUTER MEMBRANE. The rigid layer is called PEPTIDOGLYCAN and is the SHAPE-FORMING component of the cell. The outer membrane also functions as a SELECTIVE GATE but its selectivity is much LESS specific than that of the cytoplasmic membrane. The outer layer of the outer membrane in Gram negative cells is composed of a molecule called LPS. This molecule is medically significant as it is HIGHLY TOXIC to humans and death from infections of G- bacteria are often the result of poisoning by the LPS. $   % $   ( =   !   !  !    !   $ $ $!$ $ $ $!$ &$ $ $!$ $ $ $!$ 7$ $ $!$ $ $ ( ( , , ,!, , , ,!, ,), , , ,!, , , ,!, , , ,!, , , 0 0 4 4 4!4 4 4 4!4 #4 4 4!4 :4 4 4$4 4 4 4$4 4 4 4$4 ?4 4 4$4 T4 4 88 C. "G:N    4 ! R ! !6   !0 !0;K!0CPH < 0޽h ? ̙330 g_D(  D^ DS    Y Dc $ @   M Figure 7. Muramic acid subunit of the peptidoglycan of Escherichia. coli. This is the type of peptidoglycan found in most Gram-negative bacteria. The glycan backbone is a repeat polymer of two amino sugars, N-acetylglucosamine (GlNAc) and N-acetylmuramic acid (MurNAc). Attached to the MurNAc is a tetrapeptide consisting of L-ala-D-glu-DAP-D-ala. In the polymeric form of the molecule, nearby tetrapeptide side chains may be linked to one another by an interpeptide bond between DAP on one chain and D-ala on the other. Peptidoglycan structure and arrangement in E. coli is representative of all Enterobacteriaceae, and many other Gram-negative bacteria, as well. The glycan backbone is made up of alternating molecules of N-acetylglucosamine(GlNAc) and N-acetylmuramic acid (MurNAc) connected by a beta1,4-glycoside bond. The 3-carbon of MurNAc is substituted with a lactyl ether group derived from pyruvate. The lactyl ether connects the glycan backbone to a peptide side chain that contains L-alanine, (L-ala), D-glutamate (D-glu), Diaminopimelic acid (DAP), and D-alanine (D-ala). MurNAc is unique to bacterial cell walls, as is D-glu, DAP and D-ala. The muramic acid subunit shown in the Figure 7 below. Strands of murein are assembled in the periplasm from 10 muramic acid subunits, and the strands are connected to form a continuous glycan molecule that encompasses the cell. Wherever their proximity allows it, the tetrapeptide chains that project from the glycan can be cross-linked by an interpeptide bond between a free amino group on DAP and a free carboxy group on a nearby D-ala. The assembly of peptidoglycan on the outside of the plasma membrane, is mediated by a group of periplasmic enzymes which are transglycosylases, transpeptidases and carboxypeptidases. The mechanism of action of penicillin and related beta-lactam antibiotics is to block transpeptidase and carboxypeptidase enzymes in their assembly of the murein cell wall. Hence the beta lactam antibiotic are said to "block cell wall synthesis" in the Bacteria. The backbone of the peptidoglycan molecule can be cleaved by an enzyme called lysozyme that is present in animal serum, tissues and secretions, and in the phagocytic lysosome. The function of lysozyme is to lyse bacterial cells as a line of constitutive defense against bacterial pathogens. Some Gram-positive bacteria are very sensitive to lysozyme and the enzyme is quite active at low concentrations. Lachrymal secretions (tears) can be diluted 1:40,000 and retain the ability to lyse certain bacterial cells. Gram-negative bacteria are less vulnerable to attack by lysozyme because their peptidoglycan is shielded by the outer membrane. The exact site of lysozymal cleavage is the beta 1,4 bond between MurNAc and GlNAc, such that the muramic acid subunit shown in Figure 7 is the result of the action of lysozyme on bacterial peptidoglycan. "N 7`Z`Z`Z+RV1|NZ  *5  9 / 9  53 8 2 ,7 BL  2)A 9!Y, D  Q 5 &>  H D 0޽h ? ̙330 LJ(  L^ LS     Lc $O @   @The Outer Membrane of Gram-negative Bacteria Of special interest as a component of the Gram-negative cell wall is the outer membrane, a discrete bilayered structure on the outside of the peptidoglycan sheet (see Figure 9 below). For the bacterium, the outer membrane is first and foremost a permeability barrier, but primarily due to its lipopolysaccharide content, it possesses many interesting and important characteristics of Gram-negative bacteria. The outer membrane is a lipid bilayer intercalated with proteins, superficially resembling the plasma membrane. The inner face of the outer membrane is composed of phospholipids similar to the phosphoglycerides that compose the plasma membrane. The outer face of the outer membrane may contain some phospholipid, but mainly it is formed by a different type of amphiphilic molecule which is composed of lipopolysaccharide (LPS). Outer membrane proteins usually traverse the membrane and in one case, anchor the outer membrane to the underlying peptidoglycan sheet. N,It  ~~X 0 z H L 0޽h ? ̙33 0 PHT (  T^ TS    B  Tc $m @   $&e.Y|LPS lipopolysaccharide v~g Y|/f4'`̃~ހXvyr gbR _N/fYv;NT͑bR0 {^opGr024.025 Y|/fMONipQl4'`~̃~ހXgYB\vNB\S8~10nm v{|Y|{|ir(0[1u{|A08h_Y|O-yr_O NR~b0 &   ipQl4'`̃vY|bRwQ gk'` 1uN[~̃hb'}[~T S g(W̃SOˆeMbʑ>eQeg Eey:NQk }endotoxin) 0     Y|;N1u{|A08h_Y|0Tyr_'`Y|~b     {|A/fY|vk'`R0Fg̃y gqQ Tv{|A vQ;NbN:N 106D-a|S|vQ[ipQl4'`̃v{|ANdk N[hQv T k'`_N N T0 N T~̃v{|A~g g@b N T 'Y~ g7~8y [/fipQl4'`~̃Qk }vk'`-N_0     yr_'`Y|/f1uY|~bv͑ YUSMO ~g YBg zQhs(W̃SOhb SỹSObS bObS01uNTyipQl4'`̃Y|cRvz^TzzgW N T bipQl4'`̃vObS g N Tvyr_'`0ll̃^\v~̃9hncbSv N T R:N2200YOy0(WSalmonella-NY|RP[-NvO-yr_OS(uupOev@nf[elRNt[0ُ(W Oguvʋe-N g͑vaIN OY1udkS[g Oguv OgnۏL0Wt[MO0 &(WLPSv8h_Y|:STO-yr_O:S-N g Nyryrv|2-n-3-1'l|xKDO 0L-ul-D-u2^|Hep T?kS|Abq sS3 6-N1'l-D-JSsN| 0 &(A:NNy|x 1uN-YNp S|S|0xxNYyx~b [/f~̃Qk }v;NbR0TyipQl4'`̃v{|Av~gNbRg:NvQ0  K ^  $  & ad1 % *fb':    B G D   At SH t 0޽h ? ̙330 PH|(  |^ |S    MB |c $HM @  M A correlation between Gram stain reaction and cell wall properties of bacteria is summarized in Table 5. The Gram stain procedure contains a "destaining" step wherein the cells are washed with an acetone-alcohol mixture. The lipid content of the Gram-negative wall probably affects the outcome of this step so that Gram-positive cells retain a primary stain while Gram-negative cells are destained. (1) A few Gram-negative bacteria are sensitive to natural penicillins. Many Gram-negative bacteria are sensitive to some type of penicillin, especially semisynthetic penicillins. Gram-negative bacteria, including E. coli, can be made sensitive to penicillin by procedures that disrupt the permeability characteristics of the outer membrane. (2) Gram-negative bacteria are sensitive to lysozyme if pretreated by some procedure that removes the outer membrane and exposes the peptidoglycan directly to the enzyme. `Zx`Z`Z  < R& PH | 0޽h ? ̙330 TL(  ^ S    MF c $0s @  M n~ހXvR a) V[~ހYb_ b) OSRkЏR c) Ob~ހMQSYRv_c$OOYipQl~̃Sb_15~25*N'YlSvnS ipQl4'`~̃:N5~10*N'YlS) d) :Nck8^~ހRˆ@b_ e) ;b g[ir(ۏeQ~ހYipQl4'`~̃~ހXS;bRP[ϑǏ800vbu }eQ  f) N~̃vbS'`0u'`YQk } T[lṼSOvOea'`[RvsQ   7 *H  0޽h ? ̙33rRH-Xc`inruyrl3JJDh(/1>)NWb^9nt my! 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