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̙33  8(  8 8  08  8 l2 ukv[y'`$ $f3 8 c ,A3jJ?uka,$D0x 8 3 A  ?Fgruk ,$D0 8 c 6A 3jJ?Ecoli-T4 ,$D0 8 0K8   n N{|xQWb_`vuk ^bSO[yv~g(tr) e[yv~g(Fgr) YT[yv~g(Lr) 'YFg̃vTvPpelṼSO/f1u-iWb_vNASbSO4Y萌Te[yv>\~T b /fuk-N YT[yvNh0 0<}(1}50}gfgcgf cgf=cC  *H 8 0޽h ? ̙33[ S ___PPT103 +D ' b= @B D' = @BA?%,( < +O%,( < +D ' b=%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*8%(D' =-o6Bdissolve*<3<*8D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*8%(D' =-o6Bdissolve*<3<*8D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*8%(D' =-o6Bdissolve*<3<*8+   <8(  <x < c $(8  8 P < C (A 58420a x  < B8  PTobacco Mosaic Virus 0C^ < c 6A ffig41_1 H < 0޽h ? ̙33  ~@D(  Dx D c $#8  8 T D C ,A fig41_4a e D 08 0  H Adenovirus" 0 2 CR D s *iH D 0޽h ? ̙33L L(  LT L C ,Afig41_4bDrH L 0޽h ? ̙33V T(  T^ T c 6Affig41_3GH T 0޽h ? ̙33  d\\(  \x \ c $@    \ `A 3jJ ?Ecoli-T4A <$D0 P \ C (A58579a H .  \ B @17 T4 BacteriophageP0 2GOG H \ 0޽h ? ̙33aY___PPT109.+8D' b= @B D' = @BA?%,( < +O%,( < +D' b=%(%(D' =%(D3' =4@BB BB%(D' =1:Bvisible*o3>+B#style.visibility<*\%(D' =-o6Bdissolve*<3<*\+ dE(  dR d s *i d s A f?E:\virus-2-course\BIOL 230 Lecture Guide - Viral Budding.files\u2fig4.jpg@ d c $(8  8 jc$ gVvuk|P[ $3 d 0T8@`F GFig. 4: Virus Obtaining Its Envelope from Host Cell Membrane by Budding"H0 2HfH d 0޽h ? ̙33w ' l(  lR l s *. l S A ?E:\virus-2-course\BIOL 230 Lecture Guide - Electron Micrograph of HIV Budding from a Lymphocyte.files\emhivbud.jpg` l 09 CTransmission Electron Micrograph of HIV-1 Budding from a Host Cell 0D0 2BBCH l 0޽h ? ̙33+0 ++,+(  ^  S    8*  c $f @  8 "*`xQWuk|P[vg uk|P[virion sSukSO bqv0~g[tevUS*Nuk0Syuk|virus particle  8h_core 8hxMONuk|P[v-N_ sS:N8h_bWV~genome  cXcapsid ˆ}v(SV(W8h_hTV sS:NcX0/fuk|P[v;N~g/egTbSbR [8hx gOb\O(u cX|capsomerebcapsomer gbcXvN*N*Nˆ}v(| 8hcXnucleocapsid /fNUOuk@b_{wQYvW,g~g 8h_TcXTwegy:N8hcX0 Senvelope  gN~g YBgvuk vQ8hcXYNf g{|bˆ}v~bvYc y:NS0 geS N؏ g:Rzspike I{D^\ir0S[E Neg[;N~ހ FOuk9e bwQ gvQryrbSyr'`vr~g EefYNI{nBR4xOW0 V  33 uk|P[v!j_g uk|P[v[ySO6R 8^S gNASbSO[yTe[y0lṼSO-NX[(W@w YT[y0 h  34 ukv[ySO6R ukvSOb_` SmSOinclusion body agukv[;N~ހQ Qs(WIQf[>f_\ NSv'Y\0b_`0peϑ NI{v\SO y:NSmSO0(W[;N~ހQb_bSmSO/fukvyr_ N TvukvQb_bvSmSOwQ g N Tvb_`0~g0Tyr'` S(uNR{|t[0 lṼeplaque \ϑlṼSON'Yϑ[;N~ހmTT (Ws^g NmTW{Q ~Ǐpe\eb10YO\eT (W^ns^gv[;ÑԂ N g\N0 >\ g>\95nm 144*NRP[ϑ:N55,000vcX| ~b24se 0>\{9558nm -N.YT[S2.5~3.5nm _N1u24se~b 0Wg Ts -Nzz 30.5nm 0:RzT>\N@wuNs N TmQ9h :Rz20nm g8TDR>\N14052nm bbI{v$Nk 1u$Ny'Yvˆ}v(TVy\vˆ}v(gb [NN0W8TDNOea[;N~ހhbvv^SSO N 0 V ukv8hx 8hx/fuk|P[-Ng͑vbR /fukW OOo`v}SOT OSO /fuku}T;mRv;Nir(W@x0uk8hxv{|W_Y gssDNA0dsDNA0ssRNA0dsRNA0ukv8hx g~r0_N gsr g핯sb:SsI{ N T{|W0 gvuk+T g NbkN*NWV~0 h  38 r^Nh'`ukv8hx{|W NNlṼSO:NO N~ukvryrA~ke_0 iiruk0RirukvA~keu0 S8huirvuk  lṼSO N,N~lṼSObacteriophage phage ^lX[(WN6qLu-N N(W~'YYpeS8huir-NSsNv^vlṼSO0ncbS N]\OǏ5u\‰[vlṼSO\ g2850y*h vQ-N2700y/f g>\v0ncBradley1967 R_~ lṼSOqQ gmQyb_` sS C. Viral Structure Since viruses are not cells, they are structurally much more simple than bacteria. An intact infectious viral particle is called a virion (def) and consists of: 1. A genome (def) The viral genome is a single or segmented, circular or linear molecule of nucleic acid functioning as the genetic material of the virus. It can be single-stranded or double-stranded DNA or RNA (but never both), and codes for the synthesis of viral components and viral enzymes for replication. 2. A capsid The capsid (def), or core, is a protein shell surrounding the genome and is usually composed of protein subunits called capsomeres (def). The capsid serves to protect and introduce the genome into host cells. Some viruses consist of no more than a genome surrounded by a capsid and are called nucleocapsid or naked viruses (def) (see Fig. 2A and Fig. 2B and Fig. 1E). Attachment proteins project out from the capsid and bind the virus to susceptible host cells. 3. An envelope Most animal viruses also have an envelope surrounding a polyhedral or helical nucleocapsid, in which case they are called enveloped viruses (see Fig. 2C, Fig. 2D, Fig. 3, Fig. 3A, and Fig. 1H). The envelope is composed of phospholipids and glycoprotein and for most viruses, is derived from host cell membranes by a process called budding (see Fig. 4 and Fig. 3B). The envelope may come from the host cell's nuclear membrane, vacuolar membranes (packaged by the Golgi apparatus), or outer cytoplasmic membrane. Although the envelope is usually of host cell origin, the virus does incorporate proteins of its own, often appearing as glycoprotein (def) spikes, into the envelope (see Fig. 1F and Fig. 3D). These glycoprotein spikes function in attaching the virus to receptors on susceptible host cells.  V   e _ (!(! P<& U: 0  " /e     /  "   vz N -g z  * 2  W b   a!0@E!0di!0!0" ' !0 !0 !0 !0 !0  !0  !0 & !0( / !05 = !0 !0 !0  !0' . !03 :  !0@E !0di !0 !0" '  !0  !0  !0  !0   !0    !0    !0 &   !0( /   !05 =  !0  !0  !0   !0' .  !03 : H  0޽h ? ̙33T0  ,(  ,^ , S    8 , c $8 @  8 "h(uman) i(mmunodeficiency) v(irus)##>H , 0޽h ? ̙33T0  4(  4^ 4 S    8 4 c $8 @  8 "h(uman) i(mmunodeficiency) v(irus)##>H 4 0޽h ? ̙33 0 h`0@(  @^ @ S    8Z @ c $ @  8 FIGURE 41-1 The helical structure of the rigid tobacco mosaic virus rod. About 5 percent of the length of the virion is depicted. Individual 17,400-Da protein subunits (protomers) assemble in a helix with an axial repeat of 6.9 nm (49 subunits per three turns). Each turn contains a nonintegral number of subunits (16-1/3), producing a pitch of 2.3 nm. The RNA (2x1O6 Da) is sandwiched internally between adjacent turns of capsid protein, forming a RNA helix of the same pitch, 8 nm in diameter, that extends the length of virus, with three nucleotide bases in contact with each subunit. Some 2,130 protomers per virion cover and protect the RNA. The complete virus is 300 nm long and 18 nm in diameter with a hollow cylindrical core 4 nm in diameter. (From Mattern CFT: Symmetry in virus architecture. In Nayak DP (ed): Molecular Biology of Animal Viruses. Marcel Dekker, New York, 1977, as modified from Caspar DLD: Adv Protein Chem, 18:37,1963, with permission.) e[yvNhpISukTMV V  35 pISukvb_` Helical Symmetry In the replication of viruses with helical symmetry, identical protein subunits (protomers) self-assemble into a helical array surrounding the nucleic acid, which follows a similar spiral path. Such nucleocapsids form rigid, highly elongated rods or flexible filaments; in either case, details of the capsid structure are often discernible by electron microscopy. In addition to classification as flexible or rigid and as naked or enveloped, helical nucleocapsids are characterized by length, width, pitch of the helix, and number of protomers per helical turn. The most extensively studied helical virus is tobacco mosaic virus (Fig. 41-1). Many important structural features of this plant virus have been detected by x-ray diffraction studies. Figure 41-2 shows Sendai virus, an enveloped virus with helical nucleocapsid symmetry, a member of the paramyxovirus family (see Ch. 30). 6tH   w m  h I4 (5"c lXF   H @ 0޽h ? ̙33  0 t l PH (  H^ H S    8f  H c $  @  8 Icosahedral Symmetry An icosahedron is a polyhedron having 20 equilateral triangular faces and 12 vertices (Fig. 41-3). Lines through opposite vertices define axes of fivefold rotational symmetry: all structural features of the polyhedron repeat five times within each 360 of rotation about any of the fivefold axes. Lines through the centers of opposite triangular faces form axes of threefold rotational symmetry; twofold rotational symmetry axes are formed by lines through midpoints of opposite edges. An icosaheron (polyhedral or spherical) with fivefold, threefold, and twofold axes of rotational symmetry (Fig. 41-3) is defined as having 532 symmetry (read as 5,3,2). NASbSO[yvNhzukAdenovirus V  36 zukv!j_g FIGURE 41-4a Adenovirus after negative stain electron microscopy. The capsid reveals the typical isometric shell made up from 20 equilateral triangular faces. The 252 capsomeres, 12 pentons and the 240 hollow hexon capsomeres are arranged in a T = 25 symmetry pattern vetite (x 400,000). 6  B      EZ *H H 0޽h ? ̙33 0 ME`P(  P^ P S    8? P c $ @  8 }FIGURE 41-4b Adenovirus model. Capsomeres are depicted as circles surrounded by an electron dense stain. The inclined axes, h and k, are indicated. The second vertex has indices h = 5, k = 0. The total number of capsomeres C = 10(h2 + hk + k2) + 2 = 252. Capsomere organization is also expressed by the triangulation number, T, the number of unit triangles on each of the 20 faces of the icosahedron. A unit triangle is formed by lines joining the centers of three adjacent capsomeres. T = (h2 + hk + k2) = 25 for adenoviruses, and C = 1OT + 2. The 12 vertex capsomeres are surrounded by 5 other capsomeres each, therefore called penton and show 5-fold rotational symmetry. The penton base consists of 5 identical 85 kD polypeptide chains and extrudes a long antenna-like fiber protein. The 240 hexon capsomeres are trimers of the 120 kD hexon protomere polypeptide (for details see Ch. 67). ~`    { J  <  ) K )H P 0޽h ? ̙33L  0   pX (  X^ X S    8 X c $ @  8 FIGURE 41-3 Icosahedral models seen, left to right, on fivefold, threefold, and twofold axes of rotational symmetry. These axes are perpendicular to the plane of the page and pass through the centers of each figure. Both polyhedral (upper) and spherical (lower) forms are represented by different virus families. %In the adenovirus model in Figure 41-4, one of the penton capsomeres is arbitrarily assigned the indices h = 0, k = 0 (origin), where h and k are the indicated axes of the inclined (60) net of capsomeres. The net axes are formed by lines of the closest-packed neighboring capsomeres. In adenoviruses, the h and k axes also coincide with the edges of the triangular faces. Any second neighboring vertex capsomere has indices h = 5, k = 0 (or h = 0, k = 5). The capsomere number (C) can be determined to be 252 from the h and k indices and the equation: C = 10(h2 +hk + k2) + 2. 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