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H  0޽h ? ̙33 @8P(    S     l3.s^g̃=pel( $    0P04 :&b/gBl7hTEQRmS d\Oq~_15~20min[bd\O %NPls^gevW{QW)n^ &(uV-N)n0}Y'lT|Q'`S'l0(W%{Q6(     6Y Pp   d 6.r^͑l(3$3   0^@M Rl\N[ϑṽm-NṽSOǏy_bǏnRyQeg,6qTpr^(r^q)n^SǑ(u105!0100!b80!)0y͑0N,r^͑:Nn͑v10% 20% N*N~̃~ހN,͑~10-12 10-13g0 lT̃Smؚv7hT0 >NO'YFg̃N*N~ހN,͑~10 12~10 13g mSOW{Qir-N~ހSm^0R2109*N/mle 100mlW{QirS_10~90mgr^͑v~ހ0 $k0Z2K8x 2R'/'/' (( ((  (  [              B  s *޽h ? ̙33 ^V(    c ,Ao?kJml,$D0  s *` P   ^ 7.kJml"$ |  0$ St/f(WN[VQ ̃`m-Nv~ހSm^NmJm^bckk sSNIQ[^bckk ̃peY IQ[^'Y0Vdk PRNRIQIQ^ (WN[l NKm[̃`mvIQ[^ 1\SS^Q̃mvSm^02W 2V%V    0,  pyrp_0{OFOfSr^pb0  2' B  s *޽h ? ̙33 ~v(     6    8.utchl0&$3'$3(     0t`5 l`Km+T.lϑ ˆ}v(/f~ހv;Nir( +Tϑ3z[ .l/fˆ}v(v;NbR ǏKm+T.lϑ1\Scw_uirvSm^0 N,~̃+T.lϑ:Nr^͑v12.5% uk̃:N7.5% ̃:N6.0% 9hncN[SOyW{Qm-Nv+T.lϑQXNN6.25 1\SKm_|ˆ}vv+Tϑ0 vQNel +Tx0x0DNA0RNA0'lϑ0m^irϑ0NN'lSx0Nx0Np0|^I{ S(uNuϑvKm[0  2s 2 23 2 2%fs%%f3%4  &  B  s *޽h ? ̙33%: <%4%DD$(    6d̂V 7Table 1. Some Methods used to measure bacterial growth ,86`Z 8 #F  D   "@  B|@    <т VMethod   Z  s *@    <ւ [ Application   Z  s *~@    <ۂ XComments   Z  s *@    << \Direct microscopic count  Z  s *@  !  < x4Enumeration of bacteria in milk or cellular vaccines55 5 Z   s *@  #  < r.Cannot distinguish living from nonliving cells// / Z " s *@  %   <‚ e!Viable cell count (colony counts)"" " Z $ s *@  '   <\B NEnumeration of bacteria in milk, foods, soil, water, laboratory cultures, etc.OO O Z & s *@  )   <8 t0Very sensitive if plating conditions are optimal11 1 Z ( s *@ V  +V    <|V  YTurbidity measurement  Z * s *V @ V  -V    <{V  IEstimations of large numbers of bacteria in clear liquid media and brothsJJ J Z , s *V @ V  /V "  <4V  TFast and nondestructive, but cannot detect cell densities less than 107 cells per ml*UF  U Z . s *V @ V B 1V B  <V B e!Measurement of total N or protein"" " Z 0 s *V B@ V B 3V B  <V B }9Measurement of total cell yield from very dense cultures:: : Z 2 s *V B@ V B 5V B  <TV B |8only practical application is in the research laboratory99 9 Z 4 s *V B@ B 7B  < B WMeasurement of Biochemical activity e.g. O2 uptake CO2 production, ATP production, etc.XX X Z 6 s *B@ B 9B  <4B ZMicrobiological assays  Z 8 s *B@ B ;B  <B VRequires a fixed standard to relate chemical activity to cell mass and/or cell numbersWW W Z : s *B@  =  <p XMeasurement of dry weight or wet weight of cells or volume of cells after centrifugationYY Y Z < s *@  ?  <p p,Measurement of total cell yield in cultures-- - Z > s *@  A  <p Bprobably more sensitive than total N or total protein measurementsCC C Z @ s *Z C s *h+H  0޽h ? ̙331 p"(  p p  <p    p f,{N _uirvuĉ_$ff B p s *޽h ? ̙333 XP(    0pP,$D0 (X1uN_uir~ހgvQ_\ xvzvQ*NSOuX[(W@wb/g NvV0 Tekuvi_N*N~ހSO-NT*N~ހ(W TNeۏLRˆvr` y:N Tekusynchronous growth ۏL TekRˆv~ހy:N Tek~ހ0 Tek~ހSO(WNUONe;RY(W~ހhTgv TNv |_dkb_`0uSyr__N V /f~ހf[0utf[TuirSf[I{xvzvo}YPge0 x0n 0n"&D"&A"4F   S    6,"p  0  p tN0_uirv*NSOuT Teku$&ff B  s *޽h ? ̙33F2 x(  xl x 0A' ??`   x  6D)p    p f_ Tekuvel "    x S 4A?]xx~~ Tekul ,$D 0\ x 0-p  _ Tekuvel;N g$N{| sXagN[lSbc)n^0IQ~0W{QWI{0 bNck8^~ހhTg N TvhTgSS0 bl b'`Ǐn0h^y_0irtel :g b Nq_T~ހN"0Y0nY"Y B x s *޽h ? ̙33D (    c Ar333E:\97e\Ğ_RvkNe\chengxinglunwen\pic\pic\chape 7\7-3.png    S 6p   p ,Helmstetter-Cummings l (       0:p` | StNN~̃~ހO'}'}|DN]xx~~_T[n N0 ek̃`mǏ_T[n ~ހ8TDvQ NSnn NeW{QmǏn mcnm8n~ހdSwYm1mT1\SN_0RRRRˆ Negveu~ހ sS:N TekW{Q0$f0Ze"f H  0޽h ? ̙33& zr (    c A~fE:\0u N\u\Growth of Bacterial Populations.files\synchrony.jpg`   0DpP  Figure 5. The synchronous growth of a bacterial population. By careful selection of cells that have just divided, a bacterial population can be synchronized in the bacterial cell division cycle. Synchrony can be maintained for only a few generations. z 20`Z4`Z+`Z2`Z7`Z    S Op   p xFigure .synchronous growth"`Z  H  0޽h ? ̙33:;   (       <WpMMM    p bN0_uirvSOu &$     0Zp  ,$D0 :&eR/eUS~ހ_uir;NSb~̃Tuk̃ vQSOu/fNSO-N~ހpeϑvXRegh:yv & 1uN*N~ހRˆb:N$N*N~ހvey:NNN N*NNN@bve1\/fNe'eneration time, G Ne_N1\/fSO~ހpevib'YN P@b e ge_Ny:N PXe0 2$,$$$$ $$$$$BV     "  @`   c AZfhttp://www.bact.wisc.edu/Bact102/gcurve1.jpg `0,$D0   0 Yp ,$D0 6&SVh:yv/fN*N~ހ~Ǐr^NRˆTv`Q0SVS k~ǏN*NNe ~ހpev1\XRN P HTcpeXR V y:Ncpeu ُ1\/fUS~ހSOuvyr_0lM 2$8$3$$3$3 $M    s *jP] t$1.eR/eUS~ހ_uirvSOuyr_ 2&3 B   s *޽h ? ̙33    0 <& (  <R < s *s < 3 nZhttp://www.bact.wisc.edu/Bact102/gcurve1.jpgm R < s *s < 3 nZhttp://www.bact.wisc.edu/Bact102/gcurve1.jpgm   < c AZfhttp://www.bact.wisc.edu/Bact102/gcurve1.jpg0 aV  < 0j  xBy definition, bacterial growth is cell replication  i.e., growth of the culture. Most species of bacteria replicate by binary fission, where one cell divides into 2 cells, the 2 cells into 4, the 4 into 8, etc. If this cell division occurs at a steady rate  such as when the cells have adequate nutrients and compatible growing conditions  we can plot numbers of cells vs. time such as on the graph at right. Before too long, we will need to extend the paper vertically as the population continues to double. For a culture where cells divide every 20 minutes, one cell can result in 16,777,216 (i.e., 224) cells after just 8 hours  barring nutrient depletion or other growth-altering conditions.B 2y ]    <  64j!    j r$1.eR/eUS~ހ_uirvSOuyr_&3 B < s *޽h ? ̙33Z   @D(  DR D s *d D c AZfhttp://www.bact.wisc.edu/Bact102/gcurve2.jpg  ^ D 0+d  dIf we were to convert our vertical axis to a logarithmic scale  as on the graph at right  we will not need as many sheets of graph paper, and we will find that a steady rate of growth is reflected as a straight line. (On the vertical axis, the same distance on the paper is covered with each doubling.) This type of graph paper is called semilogarithmic graph paper on which we will be plotting our class results. The numbers we plot will fall on the graph at the same place the logarithms of these numbers would fall when plotted on conventional graph paper. D2  25{2T    B D s *޽h ? ̙33R! PL(  LR L s *NN L c AZfhttp://www.bact.wisc.edu/Bact102/gcurve3.jpg P L 0:d s bThe example at right shows the type of graph we may obtain from our class data. We can plot both colony-forming units (CFUs) per ml and absorbance on the same graph, remembering that the absorbance units should also be on a logarithmic scale. Rather than "connecting the dots," we draw the best straight line among our CFU/ml plots to represent the phases of growth  lag, exponential, and the start of the maximum stationary phase. 8 2~2w  6  H L 0޽h ? ̙33n +   ` (    s A~fE:\0u N\u\Growth of Bacterial Populations.files\gtproblem.jpgyP  6Ld  `&cpeuSN(u N_h:y b = B2n B, b T t S1uՋ_ n SǏ N_{_Q \I{_$NOS[pe͑cT_ lgb = lgB + nlg2 lgb - lgB lgb - lgB lg2 0.301 p 20,  )                     6]d3  b_-N B :NwY^~ހpev b :Ncpeug*Ne;R t ev~ހpev n :NNNpe2 2                 6\d @P@ Un = 2    6Hd P@@ S= 2     S $Rd0  d Tcpeu B  s *޽h ? ̙33 f^p(    0  O,$D0 *OYNW{Qm-N_uirpev1u_Yv12 000 B ~4h  t TXR0R49 000 000 b ُ7h n  (lg4.9107 lg1.2104 0.30112Vb 2L                         0$ p  ,$D0 b&PRN n T t ؏SN{Q N TW{QagN NvNeG G  t / n (W,gO-N G 460 / 12 20min 4"y_uirvNe:N20R0(W4\eQqQA~kN12N0d 2/ $       :           c A~fE:\0u N\u\Growth of Bacterial Populations.files\gtproblem.jpg @ ,,$D 0B  s *޽h ? ̙33 L(    0b` &NeYS^~̃vus New us_ Ne usba0(W_Y_uirf[xvz-N8^8^N㉮_uirvNe0 &Ne(W N Ty_uir-NvSS_'Y Ype_uirvNe:N1~3h,6q gN_uv_uirvNe؏ N0R10min, SNN_uirvNetSSQ\ebQ)YSY TNy_uir (W N TvuagN NvQNevw_N N TFO/f (WN[agN N kNy_uirvNe/fR`[v Vdk[/f_uir̃yvN*N͑yr_0t0x2"&"K"G& "        G B  s *޽h ? ̙33  <H(  <x < c $`@    < c $h  *h LB < c $DLB < c $DT T LB < c $DppLB < c $DLB < c $DLB  < c $DLB  < c $DLB  < c $DLB  < c $DLB  < c $D  LB < c $D8 8 LB < c $D  LB < c $D  LB < c $D  LB < c $DRB < s *DjJ RB < s *DjJRB < s *DjJB < s *޽h ? K09 //_`/(  4.F  _ @|-@ ~ ]~s@  "  < M Bacterium   Z ! s *p@  $  <X  JMedium  Z # s *@ ~ &~  < ~ ]Generation Time (minutes)  Z % s *~|@  (  <l VEscherichia coli Z ' s *w@  *  < Q Glucose-salts  Z ) s *n@ ~ ,~  <~ H17 Z + s *~@  .   < fBacillus megaterium    Z - s *w@  0   < Q Sucrose-salts  Z / s *n@ ~ 2~   <~ H25 Z 1 s *~@ & 4&   <l& fStreptococcus lactis  Z 3 s *&n@ & 6&   <& HMilk  Z 5 s *&n@ ~& 8~&  <~& H26 Z 7 s *~&@ &F :&F  <,&F fStreptococcus lactis  Z 9 s *&Fw@ &F <&F  <`&F Q Lactose broth  Z ; s *&Fn@ &~F >&~F  <&~F H48 Z = s *&~F@ Ff @Ff  <Ff gStaphylococcus aureus  Z ? s *Ff~@ Ff BFf  <Ff XHeart infusion broth  Z A s *Ffq@ F~f DF~f  <F~f K27-30 Z C s *F~f@ fl  Ffl   <xfl  ]Lactobacillus acidophilus  Z E s *fl n@ fl  Hfl   <fl  HMilk  Z G s *fl q@ f~l  Jf~l   <f~l  K66-87 Z I s *f~l @ l r  Ll r   <Dl r  YRhizobium japonicum Z K s *l r @ l r  Nl r   <l r  nMannitol-salts-yeast extract  Z M s *l r s@ l ~r  Pl ~r   <ěl ~r  M344-461 Z O s *l ~r @ r x Rr x  <,țr x ^Mycobacterium tuberculosis  Z Q s *r xs@ r x Tr x  <X̛r x M Synthetic   Z S s *r xs@ r ~x Vr ~x  <Лr ~x M792-932 Z U s *r ~x~@ x Xx  <ԛx XTreponema pallidum Z W s *xw@ x Zx  <x؛x Q Rabbit testes  Z Y s *xp@ x~ \x~   <ܛx~ J1980 Z [ s *x~Z ^ s *h+ ` S 8`   WTable . Generation times for some common bacteria under optimal conditions of growth. $XW3W  B  s *޽h ? ̙33d    D(  D D  <  ` @    D c $F  & KLB D c $DLB D c $DLB D c $D  LB D c $DLB D c $D0 0 LB  D c $DP P LB  D c $D  LB  D c $D LB  D c $D@ LB  D c $D`` B D s *޽h ? % F(    0   Z G (generation time) = t(time, in minutes or hours)/n(number of generations) G = t/n G = generation time (time for the cells to divide) t = time interval in hours or minutes B = number of bacteria at the beginning of a time interval b = number of bacteria at the end of the time interval n = number of generations (number of times the cell population doubles during the time interval) b = B x 2n (This equation is an expression of growth by binary fission) G = t/n ~ 2M 1 ?  6  0XP Solve for n: n = logB + nlog2 n = logb - logB log2 n = logb - logB .301 n = 3.3 logb/B G = t/n Solve for G G = t 3.3 log b/B R 2%  >             <    S (0p P   |Calculation of Generation Time"  B  s *޽h ? ̙33 _W(    c A~fE:\0u N\u\Growth of Bacterial Populations.files\gtproblem.jpgP K  0|vp Example: What is the generation time of a bacterial population that increases from 10,000 cells to 10,000,000 cells in four hours of growth? 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" &:"&!&"&!"&*  7 B  s *޽h ? ̙33F (    c A~3E:\0u N\u\Growth of Bacterial Populations.files\chemostat.jpg   0    Figure . Schematic diagram of a chemostat, a device for the continuous culture of bacteria. The chemostat relieves the environmental conditions that restrict growth by continuously supplying nutrients to cells and removing waste substances and spent cells from the culture medium. ! 2+`Z0`Z3`Z.`Z1`Z(`ZL!   9     B  s *޽h ? ̙33 jb(    c ,A(o?R`ShV ,$D 08  S T p  d .R`ShVChemostat bbactogen >&$3 $3 &$3B       B  s *޽h ? ̙334 NF(    c ,A&Ԕ?R`JmhV0 C ,$D 0  0   +,$D0 RJi_ǏW{QWAm OW{QmJm^OcR`[vޏ~W{Qel0 StǏeW{QWAmeQv^TW{QirAmQv^eg~c̃Sm^ NS,sSJm^ NS0;NǑ(uR`JmhV S_Jm^ؚe OeW{QWvAmR_ Jm^MNO RQbaW{QWvAm0 yrpW(Ǐϑ _uirY~NgؚsۏLu v^S(WAQVQc6R N TṽSO[^FO]z YBg pt0s 23$!%$!Q$$!$  $F    + 3    6 ! p   fޏ~W{Qb/g  R`JmW{Q &$3  B  0 ` 2,$D0 PO(uV(uNuN'Yϑ̃SO0uNÑSOuvs^LvgNN"Nir YsNx0YNI{0,) 2$!$$) B  s *޽h ? ̙33 $Ed(  Vf  D #"U  < ?P  n [[:N;N"  @`  < ? P v N TusṽSO  "   @`  <i?  lNONgؚ"  @`  < i?  hR`["  @`  <i?   r gP6RuVP["  @`  <i?  xW{QWAmYc6R   "   @`  <x"i?  jR`ShV"  @`  < *i?P  luN:N;N"  @`  <h2i?P   'Yϑ̃SObÑSOb_bvs^LvNir"  @`  <\:i?   hgؚ"  @`   <6___PPT9  ~Four characteristic phases of the growth cycle are recognized. 1. Lag Phase. Immediately after inoculation of the cells into fresh medium, the population remains temporarily unchanged. Although there is no apparent cell division occurring, the cells may be growing in volume or mass, synthesizing enzymes, proteins, RNA, etc., and increasing in metabolic activity. The length of the lag phase is apparently dependent on a wide variety of factors including the size of the inoculum; time necessary to recover from physiacal damage or shock in the transfer; time required for synthesis of essential coenzymes or division factors; and time required for synthesis of new (inducible) enzymes that are necessary to metabolize the substrates present in the medium. ^ߏg yrp_Ye~ހN, NzsSۏLA~k ~̃peQNN NXR f~ zs^0dkeg~ހQvN";mRkev ~ހSOyf>fX'Y (WdkgvTg \pe~ހ_YRˆ f~eu g NGS0 NuvSV/f~ހ:NN^esXۏLte kYTbev_vv0vbgN-NN"Nir b:ONYvϑbuVP[0bcye b_c$OI{0 SV1uN~ހۏeQesX, ~ހpeveX u g@b N M gN*N^Ǐ z ۏL'Yϑ vv[v vub0 ~ހyr'`:~ހveHN"^ 8^ev *NSOSOy>fWX'Y ^esX Tbevv geS0RcySO'YR{kN0Ygcy(uv/fev~ހbeW{QW%{Q N0N[ Reg^0  2 2'"N&#C&' ?' & ' $' 9'  p  !    V   H  0޽h ? ̙33 0 jb0(  X  C    b  S 9 @T  <4___PPT9 W{QWbR cy0R T7h~bvW{QWkcy0R~b N TvW{QW-N vQ^ߏgwN ^(uaIN 1uN^ߏgvwq_T_uirvck8^uhTg (WSu]NuN-N^uNhTg OMNOYv)R(us @bN(WuN[-N;`/fl)w^ߏg0 ǑSv)wlag phase vce g `$XRcyϑ SOOR----^'`X:_ a$Ǒ(u[peugvePX̃y b$teW{QWvbR (WyP[W-NReQSuW{QWvgNbR0 c$ (uA~k_ṽy t " 2d 2"&&%E '  &*y   V H  0޽h ? ̙33 0   Pp (  X  C       S Pb @   r  2. Exponential (log) Phase. The exponential phase of growth is a pattern of balanced growth wherein all the cells are dividing regularly by binary fission, and are growing by geometric progression. The cells divide at a constant rate depending upon the composition of the growth medium and the conditions of incubation. The rate of exponential growth of a bacterial culture is expressed as generation time, also the doubling time of the bacterial population. Generation time (G) is defined as the time (t) per generation (n = number of generations). Hence, G=t/n is the equation from which calculations of generation time (below) derive. [peglogarithmic phase [pegv~̃~ހN";mRg:_ Ng_v^ۏLuTA~k ̃SOpev cQUO~peXR0[N~̃eg dkegṽpeN2n^XR,nNh~ހRˆv!kpe0 ^(uaIN  /fN"utxvzvo}YPge  XklṼSOvg[;Ñ uN-NcyvgsOy dkgvutyrp ̃SON";mÍ m%{Qir(Y us_ ̃SOpev>fWXY0 ̃SO'Y\0b_`0utyr_I{kN YNgv~ހ[tSV }Oea 1uNdkegṽykePX uN N8^(uegṽSO\O:Ñy0SY [[8^(u[pegṽy\O:N[Pgexvz_uirvN"0W OxvzbۏLgr0b_`‰[I{0 Su]N N=\ϑ^g N0RؚṽSO[^ ߘT]N N=\ϑO g[_uir NۏeQdkg sa~ހpevNQUO~peXR vQ[peNeHTv~sQ|0[N~̃eg dkegṽpeN2n^XR,nNh~ހRˆv!kpe0 ^(u/fN"utxvzvo}YPge0XklṼSOvg[;Ñ0cyvgsOy K 20 2A 2Q 21fe 0ZN$$$$$ 8&.& & &$ & $$ & $& $&1 f %"&   }   6       G  X       ) H  0޽h ? ̙33R0  `(  X  C       S q @T  <4___PPT9 `(ucysNẽd\OlS\_RyvPge,(Wẽs^ghbۏLs^LR~0Gbb_R~bvQNb__vޏ~R~ _uir~ހpeϑ\@wR~!kpevXY Q\ v^ekRce_eg YgR~[v݋ _uirNNRce ~W{QT S(Ws^ghb_0RUS̃=0 <r v2r'  r  H  0޽h ? ̙33F0 (  X  C      S  @T  <4___PPT9 THQ(uVSOW{QW6Rbẽs^g,6qT\N[zʑ^v\ϑ7hT(N,/f0.2_0.5ml)R0Rs^g N,v^(uẽstmh\̃mGWSm^0Rte*Ns^ghb,~ǏW{QTcSUS*Ñ=. /fg8^(uvel.&\ v2\%T&   *   H  0޽h ? ̙330 p((  X  C      S ܁ @T  <4___PPT9 n\_RyvPge(uẽutv4lۏLN|Rzʑ,6qTS N Tzʑm\(N,/f1ml)R+RN]qSv^QtS0R50!]Sv107 cells per ml), but samples can be concentrated by centrifugation or filtration to increase sensitivity. A variation of the direct microscopic count has been used to observe and measure growth of bacteria in natural environments. In order to detect and prove that thermophilic bacteria were growing in boiling hot springs, T.D. Brock immersed microscope slides in the springs and withdrew them periodically for microscopic observation. The bacteria in the boiling water attached to the glass slides naturally and grew as microcolonies on the surface. 2. Electronic counting chambers count numbers and measure size distribution of cells. For cells the size of bacteria the suspening medium must be very clean. Such elecronic devices are more often used to count eukaryotic cells such as blood cells. 3. Indirect viable cell counts, also called plate counts, involve plating out (spreading) a sample of a culture on a nutrient agar surface. The sample or cell suspension can be diluted in a nontoxic diluent (e.g. water or saline) before plating. If plated on a suitable medium, each viable unit grows and forms a colony. Each colony that can be counted is called a colony forming unit (cfu) and the number of cfu's is related to the viable number of bacteria in the sample. Advantages of the technique are its sensitivity (theoretically, a single cell can be detected), and it allows for inspection and positive identification of the organism counted. Disadvantages are (1) only living cells develop colonies that are counted; (2) clumps or chains of cells develop into a single colony; (3) colonies develop only from those organisms for which the cultural conditions are suitable for growth. The latter makes the technique virtually useless to characterize or count the total number of bacteria in complex microbial ecosystems such as soil or the animal rumen or gastrointestinal tract. Genetic probes can be used to demonstrate the diversity and relative abundance of procaryotes in such an environment, but many species identified by genetic techniques have so far proven unculturable. _uirSOy\,*NSOu_Km[,8^/fKm[_uirSOvuϑ.Ǐ_uiruvKm[ SN[‰0WċNW{QagN0%{Qir(I{[_uiruvq_T [‰0WS^_uiruvĉ_0 Km[elSb: pel0uirϑlTutchI{el0 Methods for Counting Bacteria Direct Measurement of Microbial Growth 1. A standard plate count (viable count) reflects the number of viable microbes and assumes that each bacterium grows into a single colony; plate counts are reported as number of colony-forming units (CFU) per ml (CFU/ml) or per g (CFU/g) of sample. 2. A plate count may be done either by the pour plate method or the spread plate method. 3. In filtration, bacteria are retained on the surface of a membrane filter and then transferred to a culture medium to grow and subsequently be counted. 4. The most probable number (MPN) method can be used for microbes that will grow in a liquid medium; it is a statistical estimation. 5. In a direct microscopic count, the microbes in a measured volume of a bacterial suspension are counted with the use of a specially designed slide. Estimating Bacterial Numbers by Indirect Methods 1. A spectrophotometer is used to determine turbidity (optical density) by measuring the amount of light that passes through a suspension of cells. Plotting optical density vs. CFU/ml yields a roughly linear relationship and, once established, can be used to estimate the concentration of a bacterial cell suspension prior to performing a plate count. 2. An indirect way of estimating bacterial numbers is measuring the metabolic activity of the population, for example, acid production or oxygen consumption. 3. For filamentous organisms such as fungi, measuring dry weight is a convenient method of growth measurement. pel 8^(uegKm[7hT-N@b+T~̃0uk̃0b[P[I{US~ހ_uirvpeϑ0pelSR:NvcpelTcpel$Ny0 >Y v2(v2 v28 v2x' '' ''$'c''P''l' ''/'/%'''''/"'''o'''/,''''' '6''F''''f''f'E'''' '''''''''''''' '''''' '''''''e''v'1'''' '''_''~' '/'''f4'B                     ]    o  9  H  0޽h ? ̙33B0 (  X  C      S 4v @   &mSOl \_Km7hTZP|Rzʑ W{QT 9hncluvgNOzʑ^TQsuvgؚzʑ^ ^(u b6qs t {Q7hTUSMOSOy-N~ހvяx(>(' FS      H  0޽h ? ̙33&0 8j(  8^ 8S     8c $t @   ` &A major advantage the bacteria have is that the vast majority of them are capable of rapid growth rates. Many bacteria can divide, that is, produce a NEW GENERATION, every 20 to 30 minutes under optimal environmental and nutrient conditions. Whereas, the human generation time is ~25 years. All forms of life are capable of what is called exponential or logarithmic growth in which the numbers of a species double each generation (e.g. 1, 2, 4, 8, 16, 32, 64 etc.). When this data is plotted as the numbers of a species vs. time it produces a classical growth curve that looks like this: &Figure 1. Typical bacterial exponential Growth Curve.In a rich culture medium bacteria, grown under aerobic conditions, achieve a final concentration of 2-5 x 109 cells per ml in about 12-18 hours. Although plotted on a different time scale the human growth curve looks the same; the human population at similar points on the growth curve are shown in red. &In the case of bacteria the timeline could, in theory, start with a single cell and end 24 hours later (assuming no death) with a bacterial-mass near the weight of the earth. It is interesting to note that the human population growth curve looks exactly the same; the population reached 6 billion people this month and it may double again by the time you are my current age. Another way to look at this is that a single bacterium can produce, in about 12 hours, ~5 billion offspring in two milliliters of medium; something it has taken the human 3.5 billion years to do. Unhindered growth, of course, requires unlimited nutrient supply and an unchanging environment, however, in real life this is impossible. At some point, usually sooner rather than later, something becomes unfavorable. Usually a critical nutrient becomes limiting. For example, the supply of the energy source may run out, or a trace element may become limiting or a predator may appear or waste will become toxic or a disease will take hold; any of these inhibit and eventually stop growth. The norm in nature is for a catastrophic crash to occur following a period of rapid, vigorous reproduction. In such cases the population usually undergoes a rapid die-off and the organism may even become extinct in a local region. The cycles of population increase, followed by a rapid population decline are typical of the natural world. One can not help but wonder if such a fate may await the human population. A major advantage of the rapid reproduction time, and one that makes the bacteria so successful, is that they are able to evolve so much faster. That is, as you will see in Chapters 6 and 8, evolution occurs through mutations that produce variations in the offspring of organisms. Then the environment determines which, if any mutations are advantageous. Those organisms with the advantageous mutations tend to survive and live on. Because of their faster generation times bacteria can test billions of mutations for survival while a single human may take 25 or more years to test a given mutation. Watch bacteria grow. In this Chapter you will see what bacteria require in order to grow. It should be no surprise to you to learn that, in general, they have the same needs we, and every other form of life, require (however, their biochemistry can get rather kinky). dh) j!lf"9p        9!0:!0 H 8 0޽h ? ̙33'0 ^V`@(  @X @ C    V @ S  @   zBy definition, bacterial growth is cell replication  i.e., growth of the culture. Most species of bacteria replicate by binary fission, where one cell divides into 2 cells, the 2 cells into 4, the 4 into 8, etc. If this cell division occurs at a steady rate  such as when the cells have adequate nutrients and compatible growing conditions  we can plot numbers of cells vs. time such as on the graph at right. Before too long, we will need to extend the paper vertically as the population continues to double. For a culture where cells divide every 20 minutes, one cell can result in 16,777,216 (i.e., 224) cells after just 8 hours  barring nutrient depletion or other growth-altering conditions. 6y^  H @ 0޽h ? ̙33(0 d\pH(  HX H C    \ H S   @   dIf we were to convert our vertical axis to a logarithmic scale  as on the graph at right  we will not need as many sheets of graph paper, and we will find that a steady rate of growth is reflected as a straight line. (On the vertical axis, the same distance on the paper is covered with each doubling.) This type of graph paper is called semilogarithmic graph paper on which we will be plotting our class results. The numbers we plot will fall on the graph at the same place the logarithms of these numbers would fall when plotted on conventional graph paper. 635{8T    H H 0޽h ? ̙33)0 bZP(  PX P C    Z P S  @   bThe example at right shows the type of graph we may obtain from our class data. We can plot both colony-forming units (CFUs) per ml and absorbance on the same graph, remembering that the absorbance units should also be on a logarithmic scale. Rather than "connecting the dots," we draw the best straight line among our CFU/ml plots to represent the phases of growth  lag, exponential, and the start of the maximum stationary phase. 6~8w  6  H P 0޽h ? ̙33+0 xp`(  `X ` C    p ` S ? @   DWhat Is a Colony-Forming Unit? When we are putting bacterial cells in contact with the medium in a petri plate  such as when we inoculate a certain amount of a natural sample into a plate or use a loop to streak a plate from a sample or culture (just to give two of many possible examples)  we expect the microscopic cells to multiply and ultimately form masses of cells visible to the naked eye. These macroscopic masses are called colonies, having been formed by cells which are able to utilize the nutrients in the medium under the conditions in which the plates are incubated. Besides nutrients, we have to consider temperature, oxygen availability, and other factors that are conducive for metabolism. As stated in the introduction to Experiment 1 in the lab manual: "When we observe colonies, we cannot assume each arose from just one cell originally planted on the medium, however. A pair, chain or cluster of cells which 'land' on the medium in close proximity to each other can multiply and produce a single colony. Thus, we use the term colony-forming unit when we consider the common origin for the cells of any colony." This term is usually abbreviated CFU. In the diagram at right (wherein we exaggerate the size of the cells as well as the amount of liquid in the tube), we have  as an example  a test tube containing one ml of a water sample which contains some bacterial cells, and we pour that one ml into a plate in order to determine something about the number of viable bacteria in that one ml sample of water. (Normally we would not know how many cells we are dealing with, nor how they are distributed on the plate that we inoculate, as we use this method to detect viable cells only by the visible colonies they ultimately produce.) Notice that in this scenario, three of the cells (#2, 3 and 4) are in a chain, and they land together; also two cells (#5 and 6) land close to each other. We also have a viable cell (#7) that is unable to metabolize at all under these conditions; perhaps it needs something that is not present in the medium, or the incubation temperature is too hot or cold, or there is some other reason. The cells able to metabolize and reproduce (usually by binary fission where 1 cell forms 2, the 2 form 4, the 4 form 8, etc.) ultimately form colonies during the incubation period. However, as you see, the common origin for the cells of any colony can be more than one cell, such as that chain of cells (#2, 3 and 4) or the two cells which "start out" very close together (#5 and 6). By finding three colonies arising from having inoculated that one ml of water, we cannot therefore say that there were three cells per ml of that water sample. We can only say that there were three colony-forming units per ml. Whether or not colonies are counted, they still arise from CFUs nonetheless. So "colony-forming unit" is not a term that is restricted only to quantitation. On occasion, cells of different species can constitute a CFU, and the resulting colony would then be considered a mixed culture. Hence, thorough mixing of the sample and good isolation methods are essential. Re-streaking isolated colonies on a non-selective medium can enhance the detection of different organisms. 2 P^c}YK<pd     O   =  H ` 0޽h ? ̙3330 (  X  C      S  @   mCalculation of Generation Time When growing exponentially by binary fission, the increase in a bacterial population is by geometric progression. If we start with one cell, when it divides, there are 2 cells in the first generation, 4 cells in the second generation, 8 cells in the third generation, and so on. The generation time is the time interval required for the cells (or population) to divide. G (generation time) = t(time, in minutes or hours)/n(number of generations) G = t/n G = generation time (time for the cells to divide) t = time interval in hours or minutes B = number of bacteria at the beginning of a time interval b = number of bacteria at the end of the time interval n = number of generations (number of times the cell population doubles during the time interval) b = B x 2n (This equation is an expression of growth by binary fission) G = t/n nm 2I Hm  H  0޽h ? ̙33 50   @\ (  X  C       S Ӽ @   ^  Synchronous Growth of Bacteria S zv196u) Studying the growth of bacterial populations in batch or continuous cultures does not permit any conclusions about the growth behavior of individual cells, because the distribution of cell size (and hence cell age) among the members of the population is completely random. Information about the growth behavior of individual bacteria can, however, be obtained by the study of synchronous cultures. Synchronized cultures must be composed of cells which are all at the same stage of the bacterial cell cycle. Measurements made on synchronized cultures are equivalent to measurements made on individual cells. A number of clever techniques have been devised to obtain bacterial populations at the same stage in the cell cycle. Some techniques involve manipulation of environmental parameters which induces the population to start or stop growth at the same point in the cell cycle, while others are physical methods for selection of cells that have just completed the process of binary fission. Theoretically, the smallest cells in a bacterial population are those that have just completed the process of cell division. Synchronous growth of a population of bacterial cells is illustrated in Figure 5. Synchronous cultures rapidly lose synchrony because not all cells in the population divide at exactly the same size, age or time. D`Y=8    5  H  0޽h ? ̙33>0 J(  X  C      S Ի @   L@tpeg/fNWWyr6Rv}sGr Nb gNpe[ by/f1mm2 ؚ0.1mm [y/f0.1mm3,te*Npe[RR:Nr^*N-N6___PPT9 @ 3. Stationary Phase. Exponential growth cannot be continued forever in a batch culture (e.g. a closed system such as a test tube or flask). Population growth is limited by one of three factors: 1. exhaustion of available nutrients; 2. accumulation of inhibitory metabolites or end products; 3. exhaustion of space, in this case called a lack of "biological space". During the stationary phase, if viable cells are being counted, it cannot be determined whether some cells are dying and an equal number of cells are dividing, or the population of cells has simply stopped growing and dividing. The stationary phase, like the lag phase, is not necessarily a period of quiescence. Bacteria that produce secondary metabolites, such as antibiotics, do so during the stationary phase of the growth cycle (Secondary metabolites are defined as metabolites produced after the active stage of growth). It si during the stationary phase that spore-forming bacteria have to induce or unmask the activity of dozens of genes that may be involved in sporulation process. 3z[g yrp`$eXkv~ހpeN~ހv{kNpeQNNvI{ _uirvusYNR`s^a W{Qir-Nv~ހpev0Rgؚ4ls^0 a$~ހRˆ^ NM _Yy/}Q+Tir Nb[v~̃_YNb[0 b$dkegv_uir_Yy/}N"Nir [NSuuNeg N,(W3z[gvTgNiry/}0Rؚ\ /fgsOv6eeg0 3z[gvwÑyTsXagN gsQ uN N8^8^Ǐee0pH6___PPT9 Z4. Death Phase. If incubation continues after the population reaches stationary phase, a death phase follows, in which the viable cell population declines. (Note, if counting by turbidimetric measurements or microoscopic counts, the death phase cannot be observed.). 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