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[p z6A Computational Model of C. elegans Vulval Development7 7( 7 $  x`>0e0e ?Rectangle 3 V [p with Rosie Bloxsom (Cambridge University) Tim Labeeuw (Cambridge University) Jasmin Fisher (MSR Cambridge) Hillel Kugler (MSR Cambridge)(   @` G[۴Pܝ7y___PPT10Y+D=' = @B ++DD    (z (  (& (  p@>0e0e?Title 1F  [ bC. elegans Nematode Worm   (  `>?,Content Placeholder 2 [ dModel organism for development 1mm long, about 1000 cells Completely transparent, can observe growthee e @` ( f>?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!yL/ drs/shapexml.xmlUo"7~kű i!D#t]/l%.!tof8(ºE ԛ\s<"Gx_M\v*[I벭PޙJh+Uqi7 ')GUWA-^je+ [\!ll F}J/)1M|ʰƭϜ=Y@k3QÇhDž}ѵIe35d>ڴzԃYo7$ίQK) 985&QI*6!HPR0Anfrs<K=@naO¡ nɪ>д%I*We̳=P^$voP*nRkwz&!sa,˷OZ uMzyh_h|tfŴ3Ly'%N2_$n|3=VurTgPżjU*[٤Н)7&6y u5dި:X턥sM:/MH簤ɒZ[Yc`;gR^T7 u@ݤHn,a^qEj-5AC][Ĉ`: ';7k ߄rh-HXAD VοO ̄3Ӂ 3MWPK!<drs/downrev.xmlDJ1B!\k)i)6c'ɐv\p[oGݳ39Y#a##L#z|@\v3Yct.C" ! ^@&vuC%uI;Zhߒ<_Z^滪,/ףT>ݢiPK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!yL/ *drs/shapexml.xmlPK-!<drs/downrev.xmlPK  WF>___PPT9   *,   ( A???$c-elegans_esaPicture 5c-elegans_esa.jpg2 G[۴Pܝ7___PPT10i.8,p +D=' = @B +" +EI    , (  ,, ,  p`>0e0e?Title 1F  [ hC. elegans VPC Differentiation   , f>?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!Cv drs/shapexml.xmlU]6}`Hm%Bh8"t0Grtxa&st0#$Y[RnECpX9O\YVR^KAQk!@YDi (ydkc0RSzIqDe)oU4} ڀKt$Oh2ą}э7D@xkټ"iU_MKrtlݥգf춾!- Dt~OR\KI|5JR] y@Jؕ r-0BOm^:m X;Fy8\as6فb?Y7:ڶ$pB|{vӫ.w|nx;/_uYs/15Wsa.wZuMzEh_$^,ɼ^2 {7x܋$`=urTgPżU)KզJн;9 &y M5d=ި&X텥sM9/MHV/ɊZ[[c`;* ( yzo0jx͐6S,a^sej AC]:t#wJ-,w6rAA ?Z4 kX G5~_f4PK!<drs/downrev.xmlDJ1B!\k)i)6c'ɐv\p[oGݳ39Y#a##L#z|@\v3Yct.C" ! ^@&vuC%uI;Zhߒ<_Z^滪,/ףT>ݢiPK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!Cv *drs/shapexml.xmlPK-!<drs/downrev.xmlPK WF>___PPT9   *,  z , RA@??Picture 2V G[۴Pܝ7___PPT10i.,BN+D=' = @B +"p+FL  _ W 0? 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EMmy.>J9Cp0]pv׏z1S,à7T1p4/|1%$YY9@1TT)#8mn Υ aPYxi*ʘ#5RotPdէIف7MC7ƍ/=[ja%g꫆&Px0¾Yu97*hYQc1+t|jJ^U鱓_ڲzެ6W7Qί={&%i"jREHC0BgQ rQW/YxKЉjs,|*9m!uͬRD7arUL= ҫN ߟvgvw]ϕ}ZռE{ V릓l<4.pIG* u2O:vN$䦟fdt{0*u]-YTe[Ei~-KݹKpmM³@lEDmVl+-eb m֑ԍWn%UÒ ە~@(?/No ʹ@<݀6x%SHj-5i V.RB($ P)|,uGf~F ^ ^WtAH-h'>X<]0]u'PK!8mEdrs/downrev.xmlDOk1GCB/&.R("Hhq3n%ɺoC{~: F1i4 [kHH֫ses9T,IQBBs4$VZ1U\9\<klLZmM踽`G_ _+}ܼ lWWEds+ƫk } '!܄~PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!5i *drs/shapexml.xmlPK-!8mE drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [5   ~y˼y˼?Rectangle 3 31  [ oA Visual Programming Language for Biological Processes This chapter presents a visual process calculus for designing and simulating computer models of biological systems. The calculus is based on a graphical variant of stochastic pi-calculus, extended with mobile compartments, and the simulation algorithm is based on standard kinetic theory of physical chemistry. The calculus forms the basis of a formal visual programming language for biology. The basic primitives of the calculus are first introduced by a series of examples involving genes and proteins. More complex features of the calculus are then illustrated by examples involving gene networks, cell differentiation, and immune system response. The main benefit of the calculus is its ability to model large systems incrementally, by directly composing simpler models of subsystems. The formal nature of the calculus also facilitates mathematical analysis of models, which in future could help provide insight into some of the underlying properties of biological systems. List of Contents: - Introduction to stochastic pi-calculus and the message-passing approach - Basic primitives: production, interaction, binding - Designing genetic oscillators in silico - A 3D representation for visualising dynamic interactions between components - A message sequence chart representation for visualising the causal execution history of a simulation - Modelling Cell differentiation in C. elegans. - Modelling MHC class I Antigen Presentation, an immune system pathway : 6   33: P(    fy˼y˼?Rectangle 7q2   n*   a   ~y˼y˼?Rectangle 3 31  [ In many respects, biological systems are like massively parallel, highly complex, error-prone computer systems. Our entire genetic code is stored in digital form as DNA, but instead of 0s and 1s there are four letters, G,A,T,C. A specialised protein travels along the DNA to read the code, like reading a sequence of instructions in a program. The code is divided into different modules, or genes. The genes produce RNA which is more accessible, like loading a module into memory. The RNA produces proteins, which fold into different shapes to perform the essential functions of a living organism. Some proteins can interact with the DNA to switch on different genes at different times, in the same way that a computer program executes different modules in response to signals from its environment. It would seem that DNA is not a single program, but more like a library of programs, which are activated by signals from their environment.      0e0e?4Slide Image Placeholder 6R   [ 33 }up](  <  f y˼y˼?Rectangle 7"#PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!4# drs/shapexml.xmlUMo6@C,l:] 8EϔDŪ)R%)N7AQ뢧A7o?JҺ?8:5Ycò3y31?J?ݼc5uMoݮK^nl)<[YRuި[BMdweE, p(qLQI6֑a07x#Iw% ̊'$HL[8,8Wά5O[)2G`kn^Sq/yd/joCnKQ&Q4ןLʃg)7Ppp)!!:+ŨEBbrH9Ai,җ!@ee(c>ѯ!Eg / HFP}*/ )Pio4n} V +9S_5w=LIaa_$wt]. Z:EԘ'5cKMY қ*=uU[VAݙVTb74!?g/$4mXBD-Y iøڃ P #*B=bZ2?dx+=@mnORsF`!2Z*Et&WԳ*01[ߠS,&BهC57/i,0['M')ظo&]Uh:|Nj<2\/?$OnSˠ%(ͯESs;s\?-YT)g_ f_ߛMZ?.OܒMOS'=H^o81/PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!4# *drs/shapexml.xmlPK-! drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   ['   ~ y˼y˼?Rectangle 3 31  [ aResearch in Systems Biology accelerated significantly around the time of the Human Genome Project. This project had the ambitious goal of mapping out the complete genetic code in humans, in order to unravel the mysteries of how the human body functions. Instead, the code raised many more questions than answers. Systems Biology is attempting to answer some of these questions, with the ultimate goal of being able to understand and predict the behaviour of biological systems. Systems Biology can be broadly divided into two complementary approaches. On the one hand, scientists are doing experiments in the lab and studying the results, in order to infer properties of biological systems. On the other, scientists are using this knowledge to build detailed models of systems on a computer and then testing these models in the lab. This is the approach that we are focussing on. Such models are a powerful tool, since they allow a biologist to simulate a range of experiments on a computer, before testing the most promising ones, saving time and resources and enabling more effective research. They can also give insight into the key mechanisms of how a biological system functions (will show an example of this later on) ZH , c   339     (  ;  fy˼y˼?Rectangle 7""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!R6e drs/shapexml.xmlUMo6 x-mci70=H;E8E.zZ8yfӱP ˍCęԉIs4>.[#Μ:h9'XN\vrwޗv%;YRc/3K.tR{qPڝ( k~P)ז,O%=δ(pLII6Ƒa87Gx#I% Ԋg$$͝EV1N7Ε3kNk hg뗗T>bR71ŵ(Ue y^;svq܉шЉDw1ko)Vznh? lJafi`Ȭ+ |=M\* \f}N/_>jj.Qd1 \QJüs~\[hk:ak<ĭY?gqv0SyT"hɢ* {^|Mԭ9pwіl[W)<)H&X W M8JӭSXRTNw[[[c`/^I rP޺X_daJIV!0$Dt3 v:D# ݒ_{fn #]DcRkaƉGC-JEvrϯͳS*^@0EUmujOSxc` R`MPK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!R6e *drs/shapexml.xmlPK-!Ed drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [   ~ y˼y˼?Rectangle 3 31  [ XAs scientists understand more and more of the molecular details of biological systems, they are publishing these details in increasingly complex models. For example, here is a model of an EGFR pathway, extensively studied in cancer research. (*Biological systems are like massively parallel, highly complex, error-prone computer systems. *) Y Y:Z c   33& (  9  fy˼y˼?Rectangle 7" PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!ﵰ drs/shapexml.xmlUn6?^[;* ]SLIT"Ur8 X=P ߼y3P)֕F'<0L~JϏބ3΅2Z&(t7멫>nZ'|}=]pL-5 c+ᱴOJ'U? (5E(kKVv_[V2+Ѣ¹2'%٘;G8V&۹7r+_ 1m,pYlqYkR^A?`=@ݾ<^Ɂ_4VTMQ0Gh4h8&B'YQ4 Gbx||1%$Y[魯@ TTW)#8mRRrT0*LU'R?<EZ(O%!#J񏒷MC7ƭ/=[a%gG \__C>,h}M0*hY Qr֚ Z,3U-JoIT/]Y=uo6[Q˿n@(7PӴbb d%* !jCs` yh9e(K}bַR~>IJani`Ȭ!|;M\3* \}Nϻ_6 eִټKԟ,`㡝tK:Py׿g׋0^ƽ8{2͆qoMn5S%n%욪̯e[p{ws\?>-YV)g_ f_ߛMZ?.OܒMOS'=H^o81/PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!ﵰ *drs/shapexml.xmlPK-!drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [   ~y˼y˼?Rectangle 3 31  [  However, as these models grow in size they are becoming increasingly difficult to understand, maintain and extend. Many published models consist of hundreds of reactions, but as our knowledge of biological systems continues to increase, models consisting of tens of thousands of reactions will soon be commonplace. Clearly, writing a model of this size as a single list of reactions is not a scalable solution, for the same reasons that we would not write a large software program as a single list of thousands of instructions. Rather, we need a way of decomposing a large biological model into a collection of smaller, more manageable building blocks. In computer science, many high-level languages have been developed to improve modularity. But biological programs are different from traditional computer programs in two fundamental ways: 1) they are massively parallel, and 2) each instruction has a certain probability of executing, so we are never quite sure what is going to happen next. This suggests the need to develop specialised programming languages for biology. (*Such models are a powerful tool, since they allow a biologist to simulate a range of experiments on a computer, before testing the most promising ones, saving time and resources and enabling more effective research. They can also give insight into the key mechanisms of how a biological system functions (will show an example of this later on)*)    33 (  :  fy˼y˼?Rectangle 7"!PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!ȗ drs/shapexml.xmlUMo6@,l:] 8EϔDŪ)R%)N7AQ뢧A7o? Һw?t8:1i#ê5yS?I?ݼc9s%ʈ/gKv)^fl!<]ZB{Ψ]\҇md%wey ,`ӵ(pLQI6Ƒa0Gx#I% Ԋ'$HL[8,w8Wέ5O;)RGak_^Rq/~ɤ/*ocfkQ&tx)nӝLɣg q <7 ŌgijTEܢR!SqX;O\YJ]KAQk"2tWHpm$t>HLO(?J^7ŷw>sdt~F#(Ӈ`ca_įwtU, Z:AԈǜcOLQ 29weSVAݙN[b4!?g$gTl,_MMϱ7 h9g_ f_۹S._ADJOinɦ/s|Ͼ \ PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!ȗ *drs/shapexml.xmlPK-! ȧ drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [`   ~@y˼y˼?Rectangle 3 31  [ DIn recent years, there has been considerable research on designing programming languages that are suitable for developing complex parallel computer systems. Interestingly, some of this research is also applicable to biological systems. One example of such a language is the stochastic pi-calculus. The pi-calculus was introduced by Turing award winner Robin Milner and colleagues. More recently, Corrado Priami developed a stochastic version. But it was Aviv Regev, Udi Shapiro and colleagues who first pioneered the use of stochastic pi-calculus for modelling biological systems. We are building on this pioneering work. *** [If we think about computer networks, there are lots of things happening in parallel on different machines. They can interact, but we're not sure when. We don t know when a user is going to switch off a machine. Don t know when a cell is going to die. How do we program such systems in a robust manner?]    33^w (     0e0e?4Slide Image Placeholder 1R   [   y˼y˼?(Notes Placeholder 2 31  [ Building on the work of Regev, Shapiro, and other. We developed a provably correct simulation algorithm for the stochastic pi-calculus, based on principles of exact simulation of chemical systems We also developed a graphical representation for the stochastic pi-calculus, to make the calculus more accessible to biologists. We proved the equivalence between the textual and graphical representations, showing that they can be used interchangeably. We used this algorithm to implement a stochastic simulator in F# (The nice thing about F# is that it is very close to our formal specification. This allowed rapid implementation, and also improved our confidence in the simulation results.) (     @y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!dAC drs/shapexml.xmlUn6?.,vUX虒X5E$)}CqEa9yfRl'+Nx~ԙK_n{SΜ:ht}gf8ݬNzl#+ޛZjVciNj/<.T8(5B([+KVv[YV qE+ת%oTZR"r1wG4#D4uA[isg_DW#R 'pO?EM/lu.ChǓ  h:%tb&ep`p|AgΞ3v(x<8a11NzG7(-:CԄGsBg_u[jUغ+z#jOm}C[i~~1o3SvO \UÈ#0Io8{"'b8Eit't{1qh3hɢ*ۦ*+[|%\/b%K*op rڬVZz@?#?izT, K**9 {+kLlW%q|n\hѓ߷/YGRHo(Lg  r PHh#A8RyWܛuto@,-/.:_ +6_ W#с WPK!]|drs/downrev.xmlDAK@a؍Ub(b-mNlݦw{|omup5@TδR#ϗB$19a#XOOfwMUH ϵUÖ,:o)km<v:˦R+顡7{pWygz˺`?cE ˷#$d,APK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!dAC *drs/shapexml.xmlPK-!]|drs/downrev.xmlPK q2 WF>___PPT9   *$   a 3380___PPT10.X-[  (     0e0e?4Slide Image Placeholder 1R   [   y˼y˼?(Notes Placeholder 2 31  [ cThe stochastic pi-calculus allows a system to be decomposed into modules which interact with each other by message-passing. On the left we see the traditional approach to biological modelling, where models are constructed by describing individual reactions. Each shape represents a protein in a particular state, and each box represents a reaction. The inbound edges represent reactants, and the outbound edges represent products Here, an active orange protein (Xp) can activate a purple protein (Y) by doing an "a" reaction, after which the purple protein is active (Yp) and the orange protein is inactive (X). The reverse reaction "d" can also occur. Active purple + orange -> active orange + purple. [Xp activates Y by sending a phosphate group (brown), X deactivates Yp by receiving a phosphate group] On the right we have the message passing approach. Instead of modelling the individual reactions of a biological system, we model its components. We do this by splitting each reaction into two complementary parts, as shown on the right. So reaction "a" becomes a reaction channel. An active orange protein sends on "a" and becomes inactive. An inactive purple protein receives on "a" and becomes active. And similarly for the reverse reaction. So here the description of the orange protein is decoupled from the description of the purple protein, by means of message passing. Why do we need complementarity? [to prevent a protein from reacting with itself. ] *** Essentially, the stochastic pi-calculus allows a system of chemical reactions to be rewritten as a collection of modular processes, which interact with each other by message-passing. Here is a comparison between chemical reactions and stochastic pi-calculus for a simple model of protein activation. For both models, the graphical representation at the top is equivalent to the textual representation at the bottom. The chemical reaction model of is constructed by listing the individual reactions in the system. An active protein X_{p} can interact with a protein Y through an a reaction, to produce a protein X and an active protein Y_{p}. The reverse reaction d can also occur. For the graphical representation, each shape represents a protein in a particular state and each box represents a reaction, with inbound edges (or arcs) from reactants and outbound edges to products. In contrast, the stochastic pi-calculus model is constructed by describing the behaviour of the individual components in the system. This is achieved by splitting the reaction a into two complementary actions, a send \ksend{a} and a receive \kreceive{a}, and similarly for the reaction d. Thus, an active protein X_{p} can send on \ksend{a} and evolve to a protein X, which can receive on \kreceive{d} and evolve to X_{p}. Similarly, a protein Y can receive on \kreceive{a} and evolve to an active protein Y_{p}, which can send on \ksend{d} and evolve to Y. For the graphical representation, each shape represents a protein in a particular state and each connected graph represents the set of possible states of a given protein, where a labelled edge represents an action that the protein can perform in order to change from one state to another. Since X_{p} can send on \ksend{a} and Y can receive on \kreceive{a}, the two proteins can interact with each other and evolve to a new state simultaneously. The pi-calculus model explicitly represents the fact that the X_{p} protein evolves to X and the Y protein evolves to Y_{p} after the interaction takes place. This contrasts with the chemical reaction model, which does not explicitly state which product comes from which reactant. Although we can guess by looking at the reactant and product names, we could equally well interpret the reactions to mean that X_{p} becomes Y_{p} and Y becomes X. Another feature of the pi-calculus model is that the description of the X protein can be given independently of that of the Y protein, allowing the behaviours of the two proteins to be modelled separately. Note that a notion of complementary send \ksend{a} and receive \kreceive{a} is needed in order to decompose the system into individual modules in this way. If we simply wrote X_{p}=a.X and Y=a.Y_{p} and allowed all the a actions to interact, there would be nothing to prevent two X_{p} proteins from interacting with each other, which should not be possible. The use of complementary actions in the model is also consistent with the physical characteristics of the system, since the protein X_{p} actually sends its phosphate group p to a receiver protein Y, which then becomes phosphorylated as Y_{p}. Conversely, the phosphorylated protein Y_{p} sends its phosphate group p back to protein X. 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EMxmy)>J9Ch8?rvUE Sy, T1p|5 ŔgebTܢR!S_9OYJ]JAQK2U1Hpjl$t>H̎(?J4ŷ>sdt~FPd0¾I^\,t1O8k̅ ~7Uz$a3U nh BD^HIi|3Zv҆q AFT0zFw1o)Vznwh? $Q0t4B0dֵNU&LJשg{AUz`c.ApYMꅲ@kRo_%xYaONSqL%,x9;ͬ3w&eϯX_pt{0*u]-YTeWEi~-KݹmɒJY\ N!6`q2\16OH+M*aISmaomɃJPR T/?/No ʹ@<݀6x%SDj-5i V.RB($ P)^/uGf~F ^ ^׌tAHh'>Y<_0]u'PK!wǵbdrs/downrev.xmlDOk1G~0B/E.EtkhEmM$Y7zhޏ\հYW-`:I΍u)ρ9Zbc4 zL^q%2Pyf"nbZґ*%(W Od:.TҶ~:lopʾ[!^FaS|-t+>'+ ܈~PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-! *drs/shapexml.xmlPK-!wǵb drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [   ~y˼y˼?Rectangle 3 31  [ The message-passing approach can help to reduce the size of a biological model and to improve modularity. For example, these two models represent the same system. Supposing active orange can activate purple, light blue and red. And so can active green and active blue. This leads to a combinatorial explosion in the number of reactions. Instead of describing each of these reactions individually, we can just say which proteins can send and receive on the same channel. This implies that all the senders can talk to all the receivers, without writing it down explicitly, resulting in a more compact model. This approach is also inspired by biological reality. If we think in terms of DNA, we know that complementary DNA sequences can interact. So rather than explicitly saying which sequence interacts with which other sequence, we simply check to see which sequences are complementary. The description is also more modular, since each connected graph represents the behaviour of a given protein, which is easy to change or remove altogether. @ a  :a    33    ` (     0e0e?4Slide Image Placeholder 1R   [E   y˼y˼?(Notes Placeholder 2 31  [ o5We can illustrate the modelling approach by considering one of the many published models of biological systems. 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Allows us to split a complicated model into more manageable parts. Each graph represents the behaviour of a protein in the system. The different nodes represent the states of the proteins. Proteins can change state by interacting with each other over shared channels. Improving modularity of biological models is an active area of research. This is just one way of tackling the complexity of biological models.   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[run demo] The simulation results show the population of protein P over time. The population of the gene G is not shown, since there is always one gene throughout the simulation. Initially there is one gene G and no protein P in the system. 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Y :޿ ~n,}8ZfbSvO \QÜcdћOW^E9?Wx/W/Wr3}>~(A/~PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!*Q *drs/shapexml.xmlPK-!?qdrs/downrev.xmlPK q2   j*   a 33^ ^V`>(  ` `  0e0e?4Slide Image Placeholder 1R   [ `  @y˼y˼?(Notes Placeholder 2 31  [ There are two possible reactions in the system, +b and -b. As with the previous example, the propensity \apparent{+b} of the +b reaction is given by the number of possible interactions on channel +b times the rate of a single interaction, written [X]\cdot[Y]\cdot\rate{+b}. After the interaction takes place, each X' protein is bound to a single Y' protein on a local channel -b, and the pair of bound proteins is written \new{-b}(X'\prl Y'). The number of possible interactions on channel -b is given by the number of pairs of bound X' and Y' proteins, written [\new{-b}(X'\prl Y')]. Therefore, the propensity \apparent{-b} of the -b reaction is given by [\new{-b}(X'\prl Y')]\mult\rate{-b}. If we were instead to use a non-binding interaction on channel b, then every X' protein would be able to interact with every other Y' protein, and the equilibrium would be 24 X' and Y' proteins. But since each X' protein can only interact with a single Y' protein, a reaction between X' and Y' is much less likely to occur, and the equilibrium is shifted to 97 X' and Y' proteins. This illustrates how the use of binding interactions can significantly alter the equilibrium of a system.   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Done by Elowitz and Leibler in 2000. Experiment inserts specific genes into bacteria, which produce proteins that  switch each other off . Here we have three genes, orange, blue and red. orange produces a protein that blocks blue, which in turn produces a protein that blocks red, which in turn produces a protein that blocks orange. Form a cycle. An additional gene produces a green fluorescent protein (GFP). Makes the bacteria glow. Repressed by the orange protein. This means that when the orange protein is produced, it blocks the production of the GFP. Put these together, and what hapens? The bacteria glow on and off. Continue glowing in their descendents. 0 P    3380___PPT10.SPĺ tlPpT(  p p  0e0e?4Slide Image Placeholder 1R   [ p   7y˼y˼?(Notes Placeholder 2 31  [ Here is the parameterised model of a gene gate. Each shape in the model is parameterised by its interaction channels. The genes G, B are parameterised by channels a and b, while the protein P is parameterised by channel b. Thus G(a,b) denotes a gene that receives on a and that produces proteins which send on b. The parameters allow networks of arbitrary complexity do be constructed from a single model of a gene gate. For instance, an autoinhibitory gate can be defined as G(b,b), i.e. a gene that receives on a and that produces proteins which send on b. A bistable network can be defined as G(a,b) | G(b,a) and a repressilator network can be defined as G(a,b) | G(b,c) | G(c,a).     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An extra gene for GFP can also be inserted to observe the glowing of the bacteria. We can reproduce the bacteria glowing on an off. But we also observe the oscillation of the other proteins in the simulation, which we cannot directly observe in live bacteria. In particular we see that GFP is high when there is no Tet protein, since GFP is inhibited by Tet. [g(a,b), x(a,b), Par, P(b), P0 Can add GFP: g(c,gfp) Can change behaviour of gene and see the resulting effect on the network: For example, we can add a self inhibitory behaviour to the gates: add ?b from g to x. Silences all of the oscillations. ] 0 P P 0 T   3380___PPT10.SPĺ5 p   x (  x x  0e0e?4Slide Image Placeholder 1R   [i x  =y˼y˼?(Notes Placeholder 2 31  [ YScreen shot of Visual SPiM in action. 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lJafi`Ȭ* |=M\* Tf}Nϻ_f>jj.Qd1 \QJø}4dЊ:5,歨{ۺv8],n%S9%"/̯y]s[wS>G[M]b.'Qu8IKWnHq$u㕦KI/ǰdv))PRߟxi&7ȅfZ .@ zzC}YY)y"Z4vP`+ Ml. di(o|>%E=n#q{/ i/F@ ƠNW wϬZxjPK!bdrs/downrev.xmlDMK1!ElV]R(V襵ަ$Kf{ygJuV,At1"Z` F{1ւ!6/ eCd97GN>gً4ZnhMCq0 h9PՔ)u֯ "x} iyy|6}"yǶl r PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!O7@ *drs/shapexml.xmlPK-!b drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [   ~`v(y˼y˼?Rectangle 3 31  [ Pi-calculus relies on message passing for interactions. Seems natural to express a trace of interactions as a message sequence chart. Chart semantics proved equivalent to standard reduction semantics of pi-calculus. We can describe the system, run simulations, and analyse the traces to determine causality of events. In short, we can debug a biological system in the same way that we debug a communication protocol!    3380___PPT10. p [   Pz (     0e0e?4Slide Image Placeholder 1R   [   @x(y˼y˼?(Notes Placeholder 2 31  [ vertical lines represent species, horizontal lines represent parallel composition vertical arrows represent delays, horizontal arrows represent communication.     `y(y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!)/7 drs/shapexml.xmlUn6?^K8* ؤXg3%QjTIʱ %i(uE9yfþVl'Ny.L~L3.2Z pwkk7mR.Zw{Xac j57wfew+˪"#δqZUdmIVJrcT;f~?7{䀈&ߺ; +1m,awDgSh XB9dHp꒳Cʯ(Lʽg9*HPe/<4gb(V>d*vK牔t6RRrT0<:哈:7Ru.RdէHiŁew}*CƍϜ=YZa%g'Pxsb4I,nQ$Z&t)ώc&M&? aM_VAݛF4k|C;___PPT9   *$   a 3380___PPT10.@,vi\S p(     0e0e?4Slide Image Placeholder 1R   [   @{(y˼y˼?(Notes Placeholder 2 31  [ This section describes how the repressilator model of [fig:Gene-Gate] can be simulated in 3D, in order to visualise the interactions between processes over time. All of the models in this section are displayed using the Network 3D softwareNetwork 3D is developed by Rich Williams of Microsoft Research.. [fig:3D-Gene-Gate] shows a 3D representation of the gene with negative control of [fig:Gene-Gate]. The main difference with the 2D representation is that each action is represented as a separate node instead of as an edge label. This gives rise to a representation that can be further simplified by hiding the node labels. The labels can then be revealed individually by clicking on the corresponding node. There are two types of nodes in the model: species nodes and action nodes. Action nodes are shown in green and are of fixed size, while species nodes are of variable colour and size, where the size of the node is proportional to the population of the species. There are also two types of edges: transition edges and interaction edges. Transition edges denote a transition from one species to another, and go from a species node to an action node, and from an action node to zero or more species nodes. For example, the \kreceive{a} action has one inbound edge from G(a,b) and one outbound edge to B(a,b), since the gene can transition from an active state to a blocked state by receiving on \kreceive{a}. The produce action has one inbound edge from G(a,b) and two outbound edges to G(a,b) and P(b), since the gene can produce a new protein in parallel with itself. The degrade action has only one inbound edge from P(b) and no outbound edges, since the protein disappears after the degrade action is executed. Interaction edges denote an interaction between two species, and go from a send action on a given channel to a corresponding receive action on the same channel. There are no interaction edges in [fig:3D-Gene-Gate], since there are no corresponding send and receive actions on the same channel.     7y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!fo drs/shapexml.xmlUn6}/ Zx}7`/t70)LIT"Ur8 X(ehgΜ~x,;H cLd~Θ3΄2Z$x7k7bv.Rw{Xne Jz7R">l%+?l,+9Ӣĕ[Ude"-(ʝQw#t" [tZ ̬xBǴȯOWu3R ͏p$O? EMrmy-9C0F9;|1Sy,C c8 |1%$YY魯@12!SqX;O\YJ]KAQkЏ*ʘ{"pjl$t>GJN(郯W4} R +9SConn z'yra "j̓XSSV¯J-C|8$lՖCPf6 Dr~OJ^KIy|5Zv҆%C3L` h9d s}b7R~$Q0t;1M6NJ&LJ7gAUz`c.Ap,&Bهc5/ i,1['M')L%],_'l8X.;`&x*o___PPT9   *$   a 3380___PPT10.+`շWT (     0e0e?4Slide Image Placeholder 1R   [   @~(y˼y˼?(Notes Placeholder 2 31  [ [fig:3D-Repressilator] shows a 3D representation of the repressilator model of [fig:Repressilator-Execution]. There are three groups of species nodes, one for each gene, represented in blue, red and orange. The species nodes at the bottom represent active and blocked genes, which have a population of 0 or 1, while the species nodes at the top represent proteins, which have variable populations. Transition edges are represented from the send action of a protein to the receive action of a corresponding gene. For example, there is an edge from the send action \ksend{c} of the blue protein to the receive action \kreceive{c} of the red gene, representing the fact that the blue protein P(c) can switch off the red gene G(c,a). [fig:3D-Repressilator-Cartoon] shows a sequence of frames from a 3D simulation of the repressilator model, based on the results of [fig:Gene-Gate]. The population of a species is visualised by dynamically altering the size of the corresponding species node. When the population is empty, the species node is obscured. The interactions between species are visualised by drawing an edge from a send action to a corresponding receive action on the same channel. When no interaction can take place, for example when there are no senders or no receivers, the interaction edge is obscured. The 3D layout allows complex interactions to be visualised while preventing edge crossings, which would have been unavoidable in 2 dimensions. The software can be used to rotate or zoom in to the model, in order to focus on a subset of interactions and species. The 3D simulation is complementary to the plots of species populations over time, since it allows the interactions between species to be visualised as the simulation progresses.     `(y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!Cc drs/shapexml.xmlUMo6 x-,'*حX虒X5E$)4(bC9yfáRl/+NxnԙK g eLQ:o3W3nV'|}=]pL-5 c+ᱴJ'U? (5A(kKVv_[V qE+7%oTZV"[r1wG49 D2uſA[isg_DWR gpO?EMr(lu)Ch+Ύ FA4:128x0Q t4 ŌgmbT%̇L~_KV(Uk#C8RZ~@)m|Ѐqk3gOV@pFXə|}}qb4c,nQ$Z&t OOc&LU қ:; uWVAݛVb4!?*y)%"jJUHƧ`=1)#F,ŃH7ρ1YzKs Fp$*Ѧm intPRD7aru< ҋsY :Οwo ~n,}8Zfb.3SvO \UÈ}>_FOp-z"MwXNv9n5档9m-YTeTee~-ۂKݻQdi[m$[QM8IK^Hs$u'M*aIS%=gaom)*PR ?/Mo ʹ@<=zx%SDj VW.᷶ mu24GW>U'E;2f#sk/ i/F@ Ơ O@u?PK!]|drs/downrev.xmlDAK@a؍Ub(b-mNlݦw{|omup5@TδR#ϗB$19a#XOOfwMUH ϵUÖ,:o)km<v:˦R+顡7{pWygz˺`?cE ˷#$d,APK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!Cc *drs/shapexml.xmlPK-!]|drs/downrev.xmlPK q2 WF>___PPT9   *$   a 3380___PPT10.+@a$  (     0e0e?4Slide Image Placeholder 1R   [   1y˼y˼?(Notes Placeholder 2 31  [ ?The nodes at the bottom represent genes, which can be in two states, on or off. The nodes at the top represent proteins, where the volume of the node is proportional to the population. The green nodes represent actions: inputs, outputs, delays. The green edges represent interactions, and go from an output to an input on the same channel. The proteins are constantly being produced and degraded. The proteins from one gene can switch off another gene. [The oscillator works by producing alternate populations of red, blue and purple proteins. These regular oscillations are used to define a notion of time. Our biological clock works using a similar principle, where a large population of a particular protein is produced every 24 hours.]    U  @1y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!87{ drs/shapexml.xmlUn6?^[c* Ǎn`SLIT"Ur8 X=P ߼y3P)֕F'<0L~JϏ^̙BB-~ k7.J{Xڧ~m *տ J o%+{د-+Od̙ݨ2졩RiZLnay"05䁨W&۹7(s+_2m-r~xB}y'h X.B9`Hp5gDŽOADŽN h2xPHh+A8R=)w?&_{XO{0ZO4 uVmj4~f=Ԣ|Ut͟PK!ˮdrs/downrev.xmlDJ@EBax7b'LkEl=3ILp˽Yӊ|hU0$ ȖN7RP|AVc,)PrtLͩJ0Ć 1v`,w{ FᦕwIJ;z0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!K4`ԯ drs/shapexml.xmlUn6?^[;*؉X虒X5E$)}CqE.zZ8yfӾRl'+NxaԙK g eLA:>SW3|ݴNzl#+>ZjVciNj/<T8\+Qj~PzWa>#δpZdMJVJdrcT;3~fQ.MuJoPV<#d[QVl#O&nx [ r2E(C¯ Lʽgh<(^j4ŔgmlT%̇LnQXZ*{6wP1>%*6k#۲:M!FWJẑ$JMTqqܝ4C{|O;x=s?ޜEml8|Ze;m)PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!K4`ԯ *drs/shapexml.xmlPK-!ԡ~drs/downrev.xmlPKq2   j*   a 3380___PPT10.ם "   m (     0e0e?4Slide Image Placeholder 1R   [   1y˼y˼?(Notes Placeholder 2 31  [ )We can refine our model of a gene to included mrna and other factors explicitly, without changing our definition of the network. Interesting effects, but similar design principles as the simple network. t=0.1, u = d = 0.0001 , t2=d2=0.01  X  1y˼y˼?6Slide Number Placeholder 3"!PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!` drs/shapexml.xmlUMo6@K?n`SLIT"Ur}CqE.zZ8yfӡTl/+yC3S1??,;cΜ:htͻ>VSW1|ݴjt+K>JjƖciNj/<*Uw]wKQh~Pz֖~b>L"Ec7$DZV"[2Cm?ny IwE) ̊'L;tT7@<ۼweԉ&d:d9?G zzCsYy)y"^´vP`+--BB[a " őRwhWl~b?HhA<tZXqP U U7PK!Mdrs/downrev.xmlDAO@&͘x1(ba DcD xC[6K)ލ=N{-z݈A& VզDwo (ja {X̯f*{6wP1>%*6kC۲:M!9u#GI2j*j{{O*;:.M|>;x{/ yuʗw/j&8~ٻZe;m)PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!` *drs/shapexml.xmlPK-!Mdrs/downrev.xmlPKq2   j*   a 3380___PPT10.:0y7# <4 (     0e0e?4Slide Image Placeholder 1R   [   1y˼y˼?(Notes Placeholder 2 31  [ We can also require proteins to form dimers or tetramers before they can inhibit a gene. In this setting, cooperativity acts to improve the regularity of oscillations by reducing the fluctuations in protein levels. In presence of cooperativity, the leaky transcription of a gene is less likely to perturb the oscillations, since at least two proteins must be produced in order to have an effect in the case of dimerization, and at least four proteins are required in the case of tetramerization. Thus, cooperativity can be seen not as an essential requirement for oscillations, but as a means of improving the stability of oscillations over a wider range of parameters.  Y  1y˼y˼?6Slide Number Placeholder 3""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!厱> drs/shapexml.xmlUM6 x-Ȼ@ˆșjTIko7. <G7o ߾;u1q&ujB?eg̙BgB-c~U k7bv.R7{Xڇne JzR o%+ۯ-+h%ݨ".iZTn`G~F?y IwE)ef#R@ͭE}: jm^3p#yd-joCnKPN&Q4IDŽNLQD(^ h2\bJHȳJs1*FbneCbrH9Ai,җ!@eUZ~ [): yQF2JSyIqHM/|ЄqkgV@pZXə|}}̇pEXؗ;] HLQc̅ ~7UzjᳰU[VAݙVTKm|CPӐnb d)* S0BgLcr^$ŬoXjs,|Gx6o~4Y:Ef()0*]UYAhg_ ߟwg_ e֤<=KÜo՟4,`S3tT0>Ga/t3z`tr=V8dQ]]h b.uv[#%K*gs t)blNZp@?#4]Fx%Lt5&+BIP4K; A.4tI╗sLJ[9nl! di(/|~)uGf~F ^ ^7tAHh'>Y<_0]uPK!~drs/downrev.xmlDN@E&䑸12#`e Dcݸ{vmJ{'.tysoYzӈ-+ą5 ϗ956I<ڞ9nJ!STPЦR"~d[3btn9I4Xs|d7:hߓ~ڙ/D>7|+Q`~0ؿ^]3􁜂m)PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!厱> *drs/shapexml.xmlPK-!~drs/downrev.xmlPKq2   j*   a 3380___PPT10.Jm$ 0 (     0e0e?4Slide Image Placeholder 1R   [`   1y˼y˼?(Notes Placeholder 2 31  [ PObserved effects of cooperativity on the Repressilator Network. The plots in row (i) show the populations of proteins P(a), P(b), P(c) over time (Ms). The plots in row (iii) show histograms of the duration of protein cycles for all three proteins. The results without cooperativity were obtained for t = 0.1, d = 0.0001, u = 0.001 and r = 1.0, while the results with tetramerization were obtained for t = 0.4, r2 = 0.0001 and r4 = 0.0001, with , d and r unchanged. Note that in presence of cooperativity the rate of gene repression can be significantly lower than in absence of cooperativity, while still observing regular protein cycles. Here repress r = 1.0 instead of 10.0 More general question: how can we design robust clocks? Clocks are important for cell division. Understanding how they work, and being able to design more robust ones.  Q W  1y˼y˼?6Slide Number Placeholder 3" PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!! drs/shapexml.xmlU]6}`B/PV+t٪N,) [KէCǓ3g?KҺGzIO1qsЙPF˘c5uMOݮKw^nl)<[YRuozawjcJˊ,d2LnUIPlD*wFe~F?y IE) ̊gL{#: jm^^2p#yd-jocnkPN&D~t)DŽNLѳhԻx)<6SBBu^Q1 s+S2D:NUԵ6 * zkI-uPdէHiى%Gɛ^r AΞo3v(p88a1Xط;] HLQcͅ ~Uznaz0۝U nh B[RZJB C拉%DԒ6P a2S'AY&Gl_#tbv͍I"U M dSᛡntzvTW]<2t 7|ٝݤ^(x &\!|d1Ir1w[r5wդ3[p6Z lt{0qh3hɢ*,Jk|\hK4U :RtfZGR7^iT"K**J {kLlWz%y|v\hⓠ?6;/瘒gRnI)Lk rr1B($ P)|WJݻԵtԯ@,-':+6_5N|?jQyaOPK!KOdrs/downrev.xmlDJ@EBax7b'6k)Xl=3IL;^,VnE7 LlJS!S>QZgZ.){29P1>#:.җ5k۱:M!FWIẕIJM5uXsy5C{{~ܧB9CoEmn6KA^߮Q9!Eh rPK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!! *drs/shapexml.xmlPK-!KOdrs/downrev.xmlPKq2   j*   a 3380___PPT10.0/%  P  (  9  f@1y˼y˼?Rectangle 7" PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!"R drs/shapexml.xmlUMo6 x-\an`)z$*VM*I9v}Cq6(b]>Cq4|cAZWgR&+c}Xu9/t&2'ǛP]v*;yҝ,{g*[ }VV:8TA7a[m,YacYE|:q%)@<*&'#hmҽka+ ~iskTZ"s%G_L&@<涼j!Q4Nsv?cSB'Y a c d6ŜgejTEܢR!SqX;O\YJ]KAQk2U\~ TuPdէIى%GɛzAgΞ3fa1X;]KN5 gXSSV¯JϝD|8&lՖCPwf6 Dr~O+) 58&QK®C0 <:Ä PL">'.f}-Z/>PI"U cNL#CfSYntzvT]<2t 7|ٍs~nR/}8ZzbUtf.\ea\9,r3VӬN<{d\ūSyT2hɢ*,J{|E\і,iu5d="j g{i* iSnt)YT0U6֘<خK%BJ/$L (AoSOl.7^21%Ϥkݒ^S l"B!NHRwhWl~b?HhA<tFXqP U U7PK!rdrs/downrev.xmlDMK1!ElʺvZJX"znf?p,IzygWva>@)jMp~= | &^,Pv4GB-BJ_6l&euB:]-1u'l!5&544h>l/11c5klDy !_d4z.UGBKPK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!"R *drs/shapexml.xmlPK-!rdrs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2   [   ~ !1y˼y˼?Rectangle 3 31  [ Taking things a step further. Some of the researchers who designed the clock inside bacteria also engineered logic gates. More elaborate gene networks: 125 from a collection of building blocs. One output: GFP Two inhibitors = inputs to the system. Does the bacteria glow when one or both of the inhibitors are present? 3 different genes, 5 possible promoters, gives rise to 125 possible different networks of 3 promoter-gene units. Digital input: IPTG neutralises the Tet protein, and aTc neutralises the Lac protein. Digital output: GFP is linked to lambda. Put all of these circuits inside bacteria, and looked at those that behaved like boolean gates.    3380___PPT10.5F &   p  (  ;  f"1y˼y˼?Rectangle 7""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!q drs/shapexml.xmlUn6?^[c+* Mn`)z$*VC*I9v8E.zZ8yfӾRl'+NxaԙK_nz1g eLA:>3W3|ݬNzl++>ZjVciNj/<T8L(5B(kKVv[[V 3-*{/3xTMyO/FdO% ܊g"ͭERN7-ΕskVk GۗLK19ƛq_\|) x4^pvHe4DqLL=0E1^ҜQ[T*d*v+牔t67RRrT0*L$7_K Z /JHFP}*/ (P퉯4n} +9S?kr2@>,h}M4*hY Qr֚K>3U-JoITˇo]Y=ug6[Q˿n@(7=Pjb d%* >jCsL` qh9e K}bַR>j <| G˜6Y7:CvV)0:[gUUAh', ߟvn7m>iyy5oSvv.Ba\9\N'&Z^ {8ƃQ<\DAЩ\7UВEUe[p{ >>-YV)X<]0]}PK!uqdrs/downrev.xmlDMk1ECxnJTJN"Ԃ='o>p I.tys9EP u^ 0ҹ.~/)0QKl&9XG3̤{_.Cmƹ+RƦ%X>F[ripIs(;%s/~Fǰ)xϷi6V| g[:OV@PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!q *drs/shapexml.xmlPK-!uq drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [q   ~;y˼y˼?Rectangle 3 31  [ UIn order to model these logic gates, we defined a gene with an additional input that allows the protein to be inhibited. So we have two levels: the gene itself can be switched off, or the protein it produces can be inhibited. The presence of an inhibitor is used as an input to the system. [run simulation both with and without inhibitor] V V V  3380___PPT10.[dqJ '    (  ;  f y˼y˼?Rectangle 7""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!C  drs/shapexml.xmlUn6 x]xmŎ,dwp#N3%Qc %iPź ͛7Ïb;i]iṭΤL^ǘpӛrйPF˘㟮𱞹cfu7׳~eY Rc0KدtR{qP_Rkһudeweet0sEceJ w~ 8 Zl:XʭxBCĴH*f2!;ވZSu[P"D9왔WK%+a!mP;JcBSGCY.tbv-I$*1m!jtRD7aru< ҋNsY :?&n7m>i~~1oSvv.Ba\1N/{bFQ%ż7qt jCJpTAKU6UY/e[s{s>>-YV)F[qiqp$IFLJ;T|z%{<\j(ǐ}vs#s,y /׿,IS`x0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!M 6 drs/shapexml.xmlUMo6 x-b_XeMq)USJRo(9N(EO<G7o ?~:u1>8:5Ybmg̙BgB-c~U k7bv.R{Xڧne JսR o%+߯-+2`gZ8A@$nH0Gʤ;Wf3 $͝EVN7-Ε3kVk 'TK2ڛR|sѠ]sv$E1Sy,C?b h2ŔgebTܢR!S_9O㴹-җ!@ee(c>ѯ!Yg / HFP}*/ )Pi4n} V +9S_47P¾I\,t1O8k̅ ~7Uz$a7U nh BD^HIi|5Zv҆ AFT0zBw1o)Vznwh? $Q0t4B0dֵNU&LJשg{AUz`c.ApYMꅲ@kRo^^[o՟4,`㡙tK:*WY׿/{r6Xt<θ?te\Q=VN.,ˢ4MAW̥qdIScn'[QM8IKwHu$u㕦[I/ǰv_()PRߟxi'7ȅfZ n@ zzCsYy)y"^´vP`+)l! di(o|~(uGf~F ^ ^7tAHho'>Y<_0]u'PK!Bkdrs/downrev.xmlDOk1GCB/f-ڭQDZq37ɒdoPޏ7[U+YW-`4LΌt!~shГ|0TNRPzߤ$nhґ*(W5M1WXPbC˾Uh:o-<޴?62L&7`Wܡd1PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!M 6 *drs/shapexml.xmlPK-!Bk drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [   ~ y˼y˼?Rectangle 3 31  [ /So far, the biological examples that have been presented have been modelled at the molecular level of genes and proteins. In this section we show how the stochastic pi-calculus can also be used to model interactions at the cellular level. We consider an example from the area of developmental biology, relating to the development of the egg-laying system of the C. elegans nematode worm. We consider the phase of development when the six Vulva Precursor Cells (VPCs) decide to adopt a primary, secondary or tertiary fate, depending on their positions with respect to an Anchor Cell (AC) and on their interactions with each other. The decision of a precursor cell to adopt a particular cell fate is an important part of the development of the organism, which will help determine whether or not the adult worm can successfully reproduce. More details about the system can be found in [Sternberg-Horvitz-1986, Sternberg-Horvitz-1989, Fisher-Piterman-Hajnal-Henzinger-2007].    33 QI 1(     0e0e?4Slide Image Placeholder 1R   [f     y˼y˼?(Notes Placeholder 2 31  [ VThe biological examples presented so far have all been modelled at the level of interacting genes and proteins. In this section we show how the SPiM calculus can also be used to model systems at the level of interacting cells. We consider an example from developmental biology, relating to the development of the egg-laying system of the C. elegans nematode. C. elegans has been used to study a number of fundamental cellular processes, and many breakthroughs relating to human physiology were first discovered in C. elegans. Examples include genetic regulation of organ development and programmed cell death, together with RNA interference - gene silencing by double-stranded RNA, both of which were awarded the Nobel prize in Physiology or Medicine in 2002 and 2006, respectively. More recently, the Nobel Prize in Chemistry was awarded for the discovery and development of the green fluorescent protein, GFP, for which a significant part of the research was carried out in C. elegans. In this section we consider the phase of development in C. elegans when six Vulval Precursor Cells (VPCs) decide to adopt a primary, secondary or tertiary fate depending on their positions with respect to an Anchor Cell (AC). The decision of a precursor cell to adopt a particular fate is an important part of the development of the organism, which will ultimately determine whether or not the adult worm can successfully lay eggs. More details about the system can be found in [Sternberg-Horvitz-1986, Sternberg-Horvitz-1989, Fisher-Piterman-Hajnal-Henzinger-2007] and a recent review is presented in [Sternberg-Wormbook-2005]. Various models of VPC differentiation have been published recently, such as [Giurumescu-PNAS-2006, Fisher-Piterman-Hajnal-Henzinger-2007, Kam-Developmental-Biology-2008] to cite a few, in which the differentiation process has been highly simplified. Here we present an even simpler model, which illustrate some of the main factors involved during development. While many biological details are missing, our approach allows further model refinements to be made so that relevant details can be included incrementally. W    y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!4 drs/shapexml.xmlUo6~/ZFvtu>$*VM*I9vﻣ8 b0 ˃r4l/Jy&L\Ow3@ )? _~چaeMʷ5[Q}pԦKs?lB9pxQ-Q5T_c(4kCV~_V)O8SPYݶu& [KV#t\0nAXx)}c0zx' v?[‘= ˡ4@('] L$FS6p2!t0rtë p9z$WohÔgcbT܈LaH9_A)T!@uP?S> #p+ Jv6&#>GJ+(X~\e؀HVG L,y<9H&q |'{z%1jʳp&u݀[MZjywPPzFUu;__"DZqG).4`~XBD-YY GJ8 GdGv3RsԖZ?JhS6ނinUỡ$nd|;ғsQ:E7<ޝ;~n.;xZvd.1-^Y0E6>wsIW#2p4D`1H`<`}͢0ZQ硥9Z2X][WZuAR.f/rؖ,mT;[턡 _6OHƟ=Dz &+zh]zn!`?/䥟ȿ\hyHoWOw^,qJHmW'0%*KT 1-+H}q___PPT9   *$   a 3380___PPT10.9,P  o(     0e0e?4Slide Image Placeholder 1R   [    y˼y˼?(Notes Placeholder 2 31^  [F>___PPT9  Some of the main events that take place during normal differentiation of VPC cells in C. elegans. (1) Initially the six cells P3.p - P8.p are undifferentiated. The anchor cell sends an inductive signal, which is strongest for the cell P6.p directly in front of it. (2) The strong inductive signal causes cell P6.p to adopt a primary fate, after which it sends a lateral signal to its immediate neighbours. (3) The lateral signal causes cells P5.p and P7.p to adopt a secondary fate. (4) Eventually, the remaining cells adopt a tertiary fate.   !1y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!`G( drs/shapexml.xmlUMo6 x]x!vUX虒X5E$)4(bb9(Cs8|C^ZWgR&+cyXv&9/t&2GǛPM]pXiմuV½7ˍ->v++^x\TuKQh~Pz֖~b>LWnTIv_lD*Fe#miss@(neҝk0 i ">] NH56?;‘_CVVg"/ HFP}*/))P(y24 hٓVr~eF}|X 'Q}u0D˄N5\x)+WzSZ>~j!{يJSuP"D9G%/$4`>XBD-Y i AR0ze29>Fw1o)Vznwh? GRmFЏ&Z %Et&WԳ*01[ߠSywkwz!Кԛs4^l|j&]UFɸw=uɲ3ͣJݑ_{vn #]EcRgka8jd}&:_{~a/PK!]|drs/downrev.xmlDAK@a؍Ub(b-mNlݦw{|omup5@TδR#ϗB$19a#XOOfwMUH ϵUÖ,:o)km<v:˦R+顡7{pWygz˺`?cE ˷#$d,APK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!`G( *drs/shapexml.xmlPK-!]|drs/downrev.xmlPK q2 WF>___PPT9   *$   a 3380___PPT10.B0]c  (     0e0e?4Slide Image Placeholder 1R   [    -1y˼y˼?(Notes Placeholder 2 31  [ =  We start by constructing a simplified model of an anchor cell, together with a generic model of an undifferentiated precursor cell that can adopt one of three fates: primary, secondary or tertiary. The cell models are given in [fig:VPC-Cells], where the graphical representation at the top is equivalent to the textual representation at the bottom. The anchor cell AC can repeatedly send on \ksend{ac}, which represents a persistent inductive signal. A precursor cell can have four possible states, representing the undifferentiated cell V, the primary cell fate V1 and the secondary and tertiary cell fates V2 and V3, respectively. The undifferentiated cell V is parameterised by its position x inside the worm, which is represented by a number between 3 and 8. Cells with successive numbers are considered to be adjacent to each other, and the cell at position 6 is considered to be directly opposite the anchor cell. The undifferentiated cell is also parameterised by a rate k, which represents the strength of its interaction with the anchor cell. Cells with a low k are considered far away and only interact weakly with the anchor cell, while cells with a high k are considered nearby and interact strongly with the anchor cell. For simplicity we assume only three levels for k: low, medium and high. Each undifferentiated cell is also characterised by its left and right neighbours, given by the channels l and r, which correspond to the lateral signals that a cell can use to communicate with its neighbours. The intuition is that each cell will interact with its left neighbour on channel l and with its right neighbour on channel r. We assume that once the precursor cells have adopted a particular fate, they no longer interact with the anchor cell. Initially, a precursor cell starts out undifferentiated as V. One of three things can then happen:1. It can receive a signal on \kreceive{ac} at rate k from the anchor cell, and adopt a primary fate V1. The primary cell will then send a persistent signal to its left and right neighbours on \ksend{l} and \ksend{r}, respectively.2. It can receive a signal from its left or right neighbour on \kreceive{l} or \kreceive{r}, respectively, and adopt a secondary fate V2. 3. In absence of stimulus it can eventually adopt a tertiary fate V3, at rate w.     /1y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK! drs/shapexml.xmlUMo6 x-l'* n`S왒X5E$YJӠ(9(Cs8|}NZW]3S1>vƜ9/t&2x_M]pXiմuF½3ˍ->v++^x\Tn׻얢 n],YneY|ș%\".iJTn`x=B9 IE(af8͝E~] H56?;‘@(' `0IE1S, .p4/|1%$YY9@12!S[:O봹-:!@eU1!p#Evpjl$t>GJ(W4n} ^ +9SC///#|X ǃNzG(-:EԘ'GsB_u[j,lՖCPf6 Dr~JKIq|7Zv҆)jC3L` h9d s}b7R,n~$Q0t1MVNJ&LJWg;AUz`c.Ap,&Bه}5/-xYcƷONStK*WY_dԟwfÛ3wf~zX,Gh2 *CGs Zʶ.V4_1s7Njڒ%M·@zlEDml+-=_xU m֑ԍ4=Dx?%L5&+BIP8K; A.4I⍗sL#R[9^l!di(|~*uGf~F @,-/.:[ +6>OD+|O3LW]PK!]|drs/downrev.xmlDAK@a؍Ub(b-mNlݦw{|omup5@TδR#ϗB$19a#XOOfwMUH ϵUÖ,:o)km<v:˦R+顡7{pWygz˺`?cE ˷#$d,APK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-! *drs/shapexml.xmlPK-!]|drs/downrev.xmlPK q2 WF>___PPT9   *$   a 3380___PPT10.,9s > 0 (     0e0e?4Slide Image Placeholder 1R   [    ,1y˼y˼?(Notes Placeholder 2 31  [  A model of the full developmental system is shown in [fig:VPC-Interactions], which consists of the anchor cell and the six VPC cells, where the graphical representation at the top is equivalent to the textual representation at the bottom. Each of the VPC cells is an instance of the generic model of a single cell described in [fig:VPC-Cells]. The generic model is instantiated with different parameters depending on the position of the cell, the strength of its interaction with the anchor cell, and the neighbouring cells with which it can communicate. Cells are assumed to communicate with their left and right neighbours on dedicated channels, where the left channel of a given cell is equal to the right channel of its left neighbour. For example, the cell V(6,s6,s7,high) at position 6 can communicate with its left neighbour on channel s6 and with its right neighbour on channel s7. It can also communicate with the anchor cell at a high rate, since it is assumed to be in close proximity to the anchor cell. Note that channels s3 and s9 in [fig:VPC-Interactions] are unused, since the cells to the left of position 3 and to the right of position 8 are not represented. The model can be further extended to represent these cells if necessary.    *1y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!3] drs/shapexml.xmlUn6?^[vUI N3%QjTIʱS4(bCQ4eho޼~xK*Sq&unJ?LJ3.2Z(xWkk7kR .Zw{Xac j5ZT o%+߯-'iQʍ :֨v̇:aaLs=BwV___PPT9   *$   a 3380___PPT10.,' G?P '(     0e0e?4Slide Image Placeholder 1R   [[    *1y˼y˼?(Notes Placeholder 2 31^  [F>___PPT9  =In order to simulate the process of VPC differentiation, we assign approximate rates to the generic VPC model of [fig:VPC-Cells] and to the overall system model of [fig:VPC-Interactions]. The rates of low, medium and high inductive signalling are given by 0.0, 1.0 and 10.0, respectively, while the rate ls of lateral signalling is given by 1.0. We assume that the rate of lateral signalling is the same for all precursor cells, with the rates of channels s4,\ldots,s8 being equal to ls. The rate w for a cell to adopt a tertiary fate in the absence of external signals is given by 0.0001. Although these rates are entirely arbitrary, they give an initial approximation for the relative speeds of the various interactions that take place during development, and can be adjusted accordingly as new insight is gained from simulation and experimentation. The simulation results of [fig:VPC-Simulation] show the total population of undifferentiated cells V and of differentiated cells V1,V2,V3 over time. Initially there are 6 undifferentiated cells, with one of the cells adopting a primary fate almost immediately. Two of the cells then adopt secondary fates in quick succession, and the remaining cells eventually adopt a tertiary fate.   '1y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!`G( drs/shapexml.xmlUMo6 x]x!vUX虒X5E$)4(bb9(Cs8|C^ZWgR&+cyXv&9/t&2GǛPM]pXiմuV½7ˍ->v++^x\TuKQh~Pz֖~b>LWnTIv_lD*Fe#miss@(neҝk0 i ">] NH56?;‘_CVVg"/ HFP}*/))P(y24 hٓVr~eF}|X 'Q}u0D˄N5\x)+WzSZ>~j!{يJSuP"D9G%/$4`>XBD-Y i AR0ze29>Fw1o)Vznwh? GRmFЏ&Z %Et&WԳ*01[ߠSywkwz!Кԛs4^l|j&]UFɸw=uɲ3ͣJݑ_{vn #]EcRgka8jd}&:_{~a/PK!]|drs/downrev.xmlDAK@a؍Ub(b-mNlݦw{|omup5@TδR#ϗB$19a#XOOfwMUH ϵUÖ,:o)km<v:˦R+顡7{pWygz˺`?cE ˷#$d,APK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!`G( *drs/shapexml.xmlPK-!]|drs/downrev.xmlPK q2 WF>___PPT9   *$   a 3380___PPT10.BZ UMp 5(     0e0e?4Slide Image Placeholder 1R   [i   @(1y˼y˼?(Notes Placeholder 2 31^  [F>___PPT9  KIn order to determine which cells adopt which fates, we plot the number of cells V,V1,V2,V3 at each of the 6 positions as shown in [fig:VPC-Geometry], where each circle represents a precursor cell. The horizontal coordinate denotes the cell position, which ranges from 3 to 8. The colour denotes the cell fate, where an undifferentiated cell V is shown in grey, and a cell that adopts a primary, secondary or tertiary fate is shown in red, blue or yellow, respectively. The plots are generated automatically during the simulation, by using the parameter x in processes V,V1,V2,V3 as the horizontal coordinate for each process. As a given cell adopts a particular fate, the process V(x,l,r,k) of each cell transitions to either V1(x,l,r), V2(x) or V3(x), and the processes at each position x are plotted accordingly. In this way, the SPiM simulation generates a movie of cell differentiation over time. We can take snapshots of the simulation to separate out the individual steps, as shown in [fig:VPC-Geometry]. For example, the first frame plots processes V(3,s3,s4,low)\prl V(4,s4,s5,low)\prl V(5,s5,s6,med)\prl V1(6,s6,s7)\prl V(7,s7,s8,med)\prl V(8,s8,s9,low), with process V1(6,s6,s7) at position 6 representing a primary fate and the remaining processes representing undetermined fates. As the simulation progresses, the processes interact with each other according to the behaviour outlined in [fig:VPC-Cells], with the processes V(7,s7,s8,med) and V(5,s5,s6,med) transitioning to V2(7) and V2(5), respectively, followed by the remaining processes transitioning to V3(4), V3(8) and V3(3). We can interpret the simulation results based on the model construction and the chosen parameters. The precursor cell at position 6 adopts a primary fate by interacting with the anchor cell at rate high=10. The average duration of this interaction is 0.1, which explains the rapid transition time observed in the simulation. Next, the primary cell at position 6 can signal to its left and right neighbours to adopt a secondary fate at rate ls=1.0. The average duration of these transitions is 1, which is 10 times slower than the initial inductive signal to the cell at position 6. In absence of external signals, the remaining cells adopt a tertiary fate at rate w=0.0001, which explains the length of time taken for these cells to differentiate. Although the rates are chosen arbitrarily, we observe that if we increase the rate w then the cells are able to adopt a tertiary fate more quickly. Since the simulation is stochastic, this means that the cells at positions 5 to 7 will have a greater chance of adopting a tertiary fate, even in the presence of inductive and lateral signals, which could potentially result in abnormal development. Note also that the cells at positions 5 and 7 receive an inductive signal at rate 1, which is 10 times weaker than the inductive signal to the cell at position 6. Although the inductive signal is much weaker, there is still a one in ten chance for a cell at position 5 or 7 to adopt a primary fate, resulting in abnormal development. Indeed, if we simulate the model several times we observe that cells 5 or 7 adopt the wrong fate roughly 10 percent of the time. Clearly the VPC model is not satisfactory, since in reality the process of VPC differentiation is extremely robust. Simply changing the ratio of the high to med inductive signals to something like 1000:1 in order to improve robustness is not acceptable either, since in reality the strength of the inductive signal does not differ by such a large amount between neighbouring cells. Thus, although the high-level model we have presented can mechanistically reproduce some of the stages of VPC differentiation, it cannot reproduce the inherent robustness of the process, and more refined models are needed.    %1y˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!`G( drs/shapexml.xmlUMo6 x]x!vUX虒X5E$)4(bb9(Cs8|C^ZWgR&+cyXv&9/t&2GǛPM]pXiմuV½7ˍ->v++^x\TuKQh~Pz֖~b>LWnTIv_lD*Fe#miss@(neҝk0 i ">] NH56?;‘_CVVg"/ HFP}*/))P(y24 hٓVr~eF}|X 'Q}u0D˄N5\x)+WzSZ>~j!{يJSuP"D9G%/$4`>XBD-Y i AR0ze29>Fw1o)Vznwh? GRmFЏ&Z %Et&WԳ*01[ߠSywkwz!Кԛs4^l|j&]UFɸw=uɲ3ͣJݑ_{vn #]EcRgka8jd}&:_{~a/PK!]|drs/downrev.xmlDAK@a؍Ub(b-mNlݦw{|omup5@TδR#ϗB$19a#XOOfwMUH ϵUÖ,:o)km<v:˦R+顡7{pWygz˺`?cE ˷#$d,APK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!`G( *drs/shapexml.xmlPK-!]|drs/downrev.xmlPK q2 WF>___PPT9   *$   a 3380___PPT10.B` ld L(     0e0e?4Slide Image Placeholder 1R   [   6hy˼y˼?(Notes Placeholder 2 31^  [F>___PPT9  b  To illustrate our modelling approach, we describe a refined version of the VPC model of [fig:VPC-Cells], based on the early results of [Sternberg-Horvitz-1989]. The refined model is presented in [fig:VPC-Sternberg]. One of the simplifying assumptions of the initial model was to represent a change in cell fate by the activation of a single process inside the cell, representative for example of the activation of a single gene. In the refined model we now represent a change in cell fate by the accumulation of a particular protein, where V1,V2,V3 represent proteins with primary, secondary and tertiary functions, respectively. As with the simplified model, an undifferentiated cell V is parameterised by its position x, the strength k of its interaction with the anchor cell, and the channels l and r on which it can communicate with its left and right neighbours. As a result, the overall model of the VPC system remains the same as in [fig:VPC-Interactions], while the behaviour of an undifferentiated cell V(x,l,r,k) is substantially refined. Instead of consisting of a single process that changes state over time, the cell V now consists of multiple processes running in parallel. The processes gV1,gV2,gV3 represent the genes for proteins V1,V2,V3, while the processes gVul and gLin12 represent the genes for proteins Vul and Lin12. The genes are executed in parallel with each other inside the cell, alongside the Muv proteins. The gene gVul produces proteins in the presence of a persistent inductive signal on channel ac, where the rate of production is determined by the parameter k. The Vul proteins then initiate a persistent communication with neighbouring cells by sending on \ksend{l} and \ksend{r}. The Vul proteins can also induce the production of proteins V1 by sending on \ksend{vul}. The proteins V1 are specific to cells that adopt a primary fate and can persistently send on \ksend{v1}, which causes the Lin12 receptors to degrade. This prevents the cell from responding to its own lateral signals on \kreceive{l} and \kreceive{r}, thereby preventing it from adopting a secondary fate. Conversely, the gene gLin12 produces active receptor proteins when it receives a signal on \kreceive{l} or \kreceive{r} from its left or right neighbours. The Lin12 receptors send on \ksend{lin12} and cause the production of proteins V2, which are specific to cells that adopt a secondary fate. The proteins V2 also cause the degradation of proteins V1 by sending on \ksend{v2}, thereby preventing the cell from adopting a primary fate. Finally, the gene gV3 is constitutively expressed at rate w and produces proteins V3 that are specific to cells with a tertiary fate. These proteins are in turn inhibited by receiving on \kreceive{vul}. Thus, there is a complicated network of interactions to ensure that the cells reliably adopt specific fates depending on the signals they receive. The interactions inside a particular cell take place on channels vul,v1,v2,v3,lin12 and muv. In order to ensure that the signals inside a given cell do not interfere with those inside neighbouring cells, the channels are restricted to each cell, represented by placing them in brackets inside the cell definition. It is this notion of restricted channels that enables boundaries between cells to be established in the model.     4hy˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!`G( drs/shapexml.xmlUMo6 x]x!vUX虒X5E$)4(bb9(Cs8|C^ZWgR&+cyXv&9/t&2GǛPM]pXiմuV½7ˍ->v++^x\TuKQh~Pz֖~b>LWnTIv_lD*Fe#miss@(neҝk0 i ">] NH56?;‘_CVVg"/ HFP}*/))P(y24 hٓVr~eF}|X 'Q}u0D˄N5\x)+WzSZ>~j!{يJSuP"D9G%/$4`>XBD-Y i AR0ze29>Fw1o)Vznwh? GRmFЏ&Z %Et&WԳ*01[ߠSywkwz!Кԛs4^l|j&]UFɸw=uɲ3ͣJݑ_{vn #]EcRgka8jd}&:_{~a/PK!]|drs/downrev.xmlDAK@a؍Ub(b-mNlݦw{|omup5@TδR#ϗB$19a#XOOfwMUH ϵUÖ,:o)km<v:˦R+顡7{pWygz˺`?cE ˷#$d,APK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!`G( *drs/shapexml.xmlPK-!]|drs/downrev.xmlPK q2 WF>___PPT9   *$   a 3380___PPT10.C0 B: "(     0e0e?4Slide Image Placeholder 1R   [V    1hy˼y˼?(Notes Placeholder 2 31^  [F>___PPT9  8  Simulation results for the VPC model of [fig:VPC-Sternberg], with low=0.01, med=2, high=10, v1=10, v2=0.1, v3=1, dV1=0.1, dV2=dV3=0.05 and all remaining rates set to 1. The results indicate the populations of processes V1, V2 and V3, which represent primary, secondary and tertiary fates, respectively, for each of the 6 cells. Both plots represent different views of the same simulation. The left plot represents a 3D view of the 18 different protein concentrations over time, 3 for each cell. The right plot gives another view of the same results, which displays the maximum populations for each of the proteins. Each cell has a dominant population of a particular protein, which corresponds to a particular cell fate. Initial simulation results for the VPC model of [fig:VPC-Sternberg] are presented in [fig:VPC-Sternberg-Simulation]. The results show the populations of processes V1, V2 and V3 for each of the 6 cells. Since these processes are parameterised by their location x, there are 18 different processes altogether. The first three lanes represent populations of V1,V2,V3 for the cell at position 3, the next three lanes represent populations of proteins for the cell at position 4, etc. We observe that the cell at position 6 adopts a primary fate, since it has a predominance of proteins V1, while the cells at positions 5 and 7 adopt a secondary fate, with a predominance of V2 proteins. Finally, the cells at positions 3,4 and 8 adopt a tertiary fate, with a predominance of V3 proteins. Rather than making a decision after receiving a single signal from a neighbouring cell, repeated signalling is required for a cell to adopt a particular fate. This improves the robustness of the system, since the decision to adopt a particular fate is now made over a period of time. An important mechanism is the amplification of a relatively small difference in inductive signal between cells, so that each cell can reliably adopt the correct fate. Some of these issues are discussed in more detail in [Giurumescu-PNAS-2006]. The main point we wish to make here is that we can start with a highly simplified model of a cell, and instantiate multiple copies of this model with different parameters to represent a system of communicating cells. We can then refine the model of a cell without changing the overall system definition. Additional details can be included in subsequent refinements, based on published results in [Sternberg-Wormbook-2005, Giurumescu-PNAS-2006, Fisher-Piterman-Hajnal-Henzinger-2007, Kam-Developmental-Biology-2008]. This highlights the main motivation for our approach, which is to construct complex systems from simpler building blocks, and to be able to refine these building blocks without having to reconstruct the entire system.     2hy˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!`G( drs/shapexml.xmlUMo6 x]x!vUX虒X5E$)4(bb9(Cs8|C^ZWgR&+cyXv&9/t&2GǛPM]pXiմuV½7ˍ->v++^x\TuKQh~Pz֖~b>LWnTIv_lD*Fe#miss@(neҝk0 i ">] NH56?;‘_CVVg"/ HFP}*/))P(y24 hٓVr~eF}|X 'Q}u0D˄N5\x)+WzSZ>~j!{يJSuP"D9G%/$4`>XBD-Y i AR0ze29>Fw1o)Vznwh? 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Aim: studying system for many years. Approached us to build a computational model. Understand the molecular mechanisms involved. Understand the dynamics of the system. Developing a programming language for modelling and simulating biological systems. Using this as a case study. The idea is that we should be able to write a program that mimics the behaviour of key parts of our immune system. Antigen processing is the means by which infections and vaccines are made ready for T cells to recognise. As T cells only recognise peptide fragments of larger protein antigens, generation, selection and presentation of these peptides plays a major role in determining whether an immune response will be effective. 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EMxmy)>J9Cp08;׏zqLT.f}-Z/c67'pT)9n#0 MSof" M^P^vXo)ywkwz!Кs4^l5.pIG* $WŤ? UԙbaGnSˠ%?VsK:pN1^|"y\ PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!߳ *drs/shapexml.xmlPK-!N drs/downrev.xmlPKq2   j*   a   r0e0e?Rectangle 2R   [(   ~Ghy˼y˼?Rectangle 3 31  [ bWe teamed up with these scientists to develop a computer model that captures their knowledge about the system. We also included some additional hypotheses We assume an equal supply of three classes of peptides with low, medium and high unbinding rates, resulting in high, medium and low affinity MHC peptide complexes. Note that we do not change the model for MHC. It is parameterised by the class of peptide. Use the high affinity peptide as the representative peptide. 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Discrimination improves the longer it waits before egressing. Able to capture high affinity peptides more effectively. Can simulate peptide editing for different values of egress. Simulation results are comparable to predictions. Also previously validated experimentally. High peptide turnover is key. So that each time a peptide is exchanged, there is an equal chance of loading a low, med or high affinity peptide. Like running the same experiment multiple times. L `0  # "Y  V $  ty˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!qޯ drs/shapexml.xmlUn6 x]x-Ŏ,bo.:EϔDŪ)R%)NJӠ(u ͛7Ïb;i]itgRg&/c~X&9/t.2w?|f8ݴNzl#+>ZjVciNj/<.T"FJ_#ޭ%+߭,+s`FE{ת%oTZR"rQ3a@4u0ſ[isgdLW#\ 폧H~29pƛe_\ ) p0_rvHU|œ S, GT/p|5 ŔgmlT%̇Ln%p#E~pjm$t>GJ(moW4n} +9S_CGQ̙d0¾I_覚EeBgh=VhTKcP-[weԽYoD-o(vK_Rjp2L,!]a| ځ P)B=bZrY0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!  drs/shapexml.xmlUMo6@C,b7N N3%QcJӠ(uA7o?+vҺ? 8:3yâsйPF˄㟮𱞺v:iﲍ`jW[ }V:R`0Wn],YneYK|q]2쾩RiJLnaGߝgf+pi`3 !"!]p5ж?Rp#}-o}asPN(Qty!W`DŽNL޳ r0_xt)!!:'٨Jy"t]͢T\ BJU~TY%<_KFV"/JHFP}*/)(P(y _2t!hٓVr~eL&C|X(¾I_覚EeBgh=VTKcP- [weԽYoD-o(vK_Rjp2_M,!]a|ځ P)B=bZrY&|*5H}_ޞUm5'kap 2PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!  *drs/shapexml.xmlPK-!|Wdrs/downrev.xmlPKq2   j*   a 33 5 o g 0 ,O (  , ,  0e0e?4Slide Image Placeholder 1R   [ ,  yy˼y˼?(Notes Placeholder 2 31  [ VCalculate peptide discrimination for different values of uT/q and egress. Note the additional filtering step. Much better discrimination in presence of tapasin. MHC is able to more effectively capture stable peptides. Simulation results are comparable to predictions. Our collaborators are currently running more quantitative experiments. ^W9z "y  U , zy˼y˼?6Slide Number Placeholder 3"PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!"Э drs/shapexml.xmlUMo6 x--_Xea=SkM*I9v8 X/9(Cs8|C^ZW331=9/t.2Gx_]pXywVV½3+>k+^x\T0J_#o%+߯-+s`gZTw\Jek%25*~wB0X#•v)܊'L;$tU?`<@xNWh X.B9`Hp4䊳cga4N ˃g$ƨ^d6^bNHȳIs1*FneCbrH9_Ais[*u)!G/ CC?>%p+EQ"/JHFP}*/))P(y _2t!hٓVr~e|Xq}Muc !jӓyBg_MZjp]غ+f+jom}C[PӔbb d%* S0RMsrAŬoXj "IJanB?b!|;M\3" Ro}NEïMeִϬZjPK!28drs/downrev.xmlDAK@FB2/b76E`j.4I 48}j0ɇY7V )jl%g Z.iw9X ؐ:.25 בes`WR{<ܴ6IRiPcG5(x(h󽧲{zOڏ7_J]D!?o6[ftzbr1D 8s\PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!"Э *drs/shapexml.xmlPK-!28drs/downrev.xmlPK q2   j*   a 336 H@P 0((  0< 0 f|y˼y˼?Rectangle 7"#PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!x. drs/shapexml.xmlUn6 x]xmG-,l&] 8EϔDj(R%)NJӠ(u ͛7Ïb;i]it‡I sйPF˄㟮𱞹cfu·׳~e[Y Rc0KЯtR{qP_R+һMdeee,8νP<(&97fo2٣p+ "!N7-ΕskVk GۗTK~29ƛq_\|) GQM.9;$|: Љ{!Nc4..|1#$Y[o9@ TTV)#8mKΥ aPUzi*R<EZ(O%!J񏒷MC7ƭϜ=Ya%g꫆a1 z'}jiTв&<5+XfZٱ_zlW7Qo={&%i"jJUH]0RQr^ŬoX} Fy$*9m!ntRD7aru< ҋN Y :? ~n,Zfb^#-~d1w \Qü}-?ǓѢ^.{qEo0_N/bjCJpTAKUyl2mAW¥,padi[ln'[>"6`q(-eb mΑԍWn%U>Ò JSU~@A@I}_ज़ ix%meKf; ABBd@7PHh+A8Ry_ܛut@,-':_ +65N|??jQyaOPK!'drs/downrev.xmlDMK@@!EFczi-ަfw{87"65&)$d SI8|3H|@$a1a`vtه*a9JCr!|QF?fur݊4 5vZmWl=P1;}߬wqI7apl9xvڡ$sPK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!x. *drs/shapexml.xmlPK-!' drs/downrev.xmlPKq2   j*   a 0  ~ y˼y˼?Rectangle 3 31  [ It turns out that the system is a bit more complicated than this. Our collaborators in Southampton suggested that the MHC complex behaves like a Venus flytrap. A peptide can enter an open MHC complex at random, a bit like a fly wandering into a Venus Flytrap (2). The complex has a special trigger mechanism, which triggers its closure depending on the shape of the peptide - much like a Venus flytrap, which has a trigger mechanism that only closes when the fly is of a certain size. Peptides that are not recognised by the MHC complex can escape (1), while peptides that are recognised as potentially harmful are captured by the MHC complex and presented at the cell surface (3). Note that the flytrap can also re-open.   0  0e0e?4Slide Image Placeholder 6R   [ 33 7   p 4 (  4; 4 f@ y˼y˼?Rectangle 7""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!@-> drs/shapexml.xmlUn6?^[c;*ĭS왒X5E$)}Cq,u ͛7b;i]it£wΤL^DŽM9s^\(eW?𾞹cfu7׳~eY Rc0KدtR{qP`ܯDBݺ^Yʲ2i4-*{/3xTMxsoߘ=Gd[_+ "kkFk GۗTK~39ƛq_\|) G('~ tJL=G0F3x&E/fAw1[o)n:| Gœ6Y5:CvV)0:[eUEAh'Y :?^ ~n̕}Zfb.Svv.Ba^5^Da4(wXDt-?oPc:U ZʶQ_ w{G1ڒm³I@lEDml+-eb mΑԍWn%U>_Ò JSU~@A@I}_ज़ ix%meKf; ABBd@wmKFX'HCqxS{R+so62 ~MPb h :@|%@|8hzE骆PK!Ndrs/downrev.xmlDMk@@e QhISWAjVfw$C87r0 ږ _Dh56Β,ws̴힮X ؐ*62 גeV8o0K=,7%ɫ4X[PaKCgn}+v]㐞Λ7ðzimw\}>?VsK:pN1^|"y\ PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!@-> *drs/shapexml.xmlPK-!N drs/downrev.xmlPKq2   j*   a 4  r0e0e?Rectangle 2R   [` 4  ~sy˼y˼?Rectangle 3 31  [ >We assume an equal supply of three classes of peptides with low, medium and high unbinding rates, resulting in high, medium and low affinity MHC peptide complexes. Note that we do not change the model for MHC. It is parameterised by the class of peptide. Use the high affinity peptide as the representative peptide. ? ? ?   33!8   8 (  8 8  0e0e?4Slide Image Placeholder 1R   [T 8  sy˼y˼?(Notes Placeholder 2 31  [ ~DRepeated open and close has similar effect to delayed egression. (Abbreviate close, open, egress) In absence of tapasin, maximum discrimination is determined by off-rate. In presence of tapasin, maximum discrimination is determined by off-rate squared Without tapasin: Better discrimination as second term tends to zero: slow close, slow egress or fast open. With tapasin: Better discrimination as second term tends to zero: decreased r, increased s, or decreased uT. Also, need to have faster o relative to egress, to ensure that complexes undergo a second filtering step.  E W 8  sy˼y˼?6Slide Number Placeholder 3" PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK! drs/shapexml.xmlUMo6 x-_Xeay=SH;E{PrEQ6eho޼~x;HJc>|7L~g eI:o> W3nQ|}]pL-5 c+ᱴJ'U? &J >l%+?l,+s`M3-*ܻUe.}SҲÎx;G!?&+pm`3 !"!]p5ж?^Rp#}-ocakPN(Q4})h0Xȣgt0P |___PPT9   *$   a 3380___PPT10.i : x p @X (  @; @ f sy˼y˼?Rectangle 7""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!w4V drs/shapexml.xmlUMo6@Cv,l:] 8EϔDŪ)R%)N7AQ뢧A7o?JҺ?8:5Ycò3y31?J?ݼc5uMoݮK^nl)<[YRuU7jmJkˊ X&akQ{ţl̻#}anFH+\K\OH/6YqYlqYkRd^A7`=@ݼ<^ɀ_ܖTM3=h<ʃg)7c z8bJHȳJs1*FbnQد'RGq, . aPYxi*ʘOzkbY8ȋB56Q:TKB$fG %oŇn[c9{:tJW ]_]]A>,(¾I^\,t1O8k̅ ~7Uz$a3U nh BD^HIi|3Zv҆q AFT0zFw1o)Vznwh? $Q0t4B0dֵNU&LJשg{AUz`c.ApYMꅲ@kRo_%xYaONSqL%,Y-gh9tIԹ<:Qֻe@ Щ\eВEUvuYע)ԝ9pߏЖ,iu5d"j g;i.iSnt+YT0U֘<خ %BJ/$L (AoSOh.7^21%O+ݒ^S l"-BB[a " őRwhWl~b?HhA<tZXqP U U7PK!'^drs/downrev.xmlDMK1@!El"MK)+ZDon%v7X%Ӊ :`>@-nmAVc,)(jy}Br` *hb )Cِ0s=Yf#rɇ,{[iP}o)j1c5k씺IR_ U4㜿^oC$91PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!w4V *drs/shapexml.xmlPK-!'^ drs/downrev.xmlPKq2   j*   a @  r0e0e?Rectangle 2R   [# @  ~@sy˼y˼?Rectangle 3 31  [ ]Used our model to explain the varying dependence of MHC on tapasin, which accounts for some of the genetic variability in the human immune system. Columns: different MHC types. Rows: with and without tapasin. Some MHC types don't work at all without Tapasin, others work equally well with and without. Currently being validated experimentally. Difficult to fuse MHC with tapasin in vitro. Our collaborators have managed to do this. Also looking at the molecular dynamics of MHC. Evidence that tapasin holds open MHC.     33 ;   Di (  D< D f sy˼y˼?Rectangle 7"#PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK! j drs/shapexml.xmlUMo6@3*؎SLIT"Ur}CqE.zZ8yfˡPl/ˍxS3)??.[cΜ:ht>SW2|ݴrnd+ >RjeciڥNj/<*T ۅ5F(ߔkKVr_[2 =δ(pLII6Ƒa03x#Iv% Ԋg$HL;8̷8WXkR^A;`=@]<^I_TTM1=gLjONw<&tb*%pwG^UL1M_L y;i.F(P-*2D:Neԥ4 *r/-Syq~5T[Um#C$D`z@1Q)]|Fи5g+C[%L}Րd8B>,~ v'~bnTв F<6+t|bRޔɩ_-zlW7Qo={!%i"jBUHC0\Q rQěŬXj3M$*9m!tRD7areN< ҫ3 :?dݸ+x\"=~d1 \QJü}9\Vo1^3l͗Ewr/Nj?cyT"hɢ* k^|E\ і,U5d"j g;i.iS4nt+ET0֘,خs%BuJ/$L (Ao]O/w^20%OV+ݐ^Q le" :D#nIQϯĽH\3A7A  .Ђx1ӵm]3%+PK!i$cdrs/downrev.xmlDOk1G~0^f)("hE6nfM$Y7zhޏXհYW-`2N΍u)G-1 `|~Z`&M8Ev o3]^B76- cxڒK}O+u\MES谹oå+b!^_yU{[l6V솻'+ ܈/PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-! j *drs/shapexml.xmlPK-!i$c drs/downrev.xmlPKq2   j*   a D  r0e0e?Rectangle 2R   [3 D  ~@sy˼y˼?Rectangle 3 31  [ mWe recently obtained a research grant from the Medical Research Council (the British equivalent of the NIH) to complete a more detailed model of our system and to experimentally test some of our hypotheses. Our modelling approach is important as we extend and modify our model.     33M u    H (  H; H f`sy˼y˼?Rectangle 7""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!  drs/shapexml.xmlUMo6@CnSLIT"Ur}CqE.zZ8yfӡTl/+yC3S1??t9/t&2Gw?|fbXYլuV}0ˍ->v++^xTwKQh~Pz֖nȀcδ(qLQI6֑a0x#Iw% ̊'$HL[8,8Wέ5O[)2G`kn^Sq/yd/joCnKQ&Q4qvUONQDb j8bFHȳJs1*FbnQد'RGqJ]JAQK2U1Hpjl$t>H̎(?J4ŷ>sdt~B[@>,(¾I^\,t1O8k̥ ~7Uz$a3U nh BD^HIi|3Zv҆q AFT0zAw1o)Vzawh?$Q0t4B0dֵNU&LJשg{AUz`c!ApyMꥲ@kRo_o՟4,f`㾙tK:*WY׿OGˤ3< ag\Fd͸+C p]AKUeQ_ +Rwn~}?B[Rx1H-v&X W M:JӭgcXRTAw[[[c`/ (O B3-O7M=,x<ǔKݑ_{vn5#]DcRgkaƉGgC-*PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!  *drs/shapexml.xmlPK-!  drs/downrev.xmlPKq2   j*   a H  r0e0e?Rectangle 2R   [ H  ~sy˼y˼?Rectangle 3 31  [ This modularity allows us to build large and complex models by incrementally composing simpler components. We can modify a particular component in a modular way, without changing the rest of the graph. The models are easier to extend and maintain. As biological models become increasingly large and complex, modular programming languages could have just as big an impact for biology as they did for computing.     33 t F > 0 L& (  L= L f`sy˼y˼?Rectangle 7"$PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!;F⺳ drs/shapexml.xmlUMo6 x-/uZ 8EϔDŪ)R%)N7AQ뢧A7o?KҺ?8:5Yb3y31?I?~j*U1y_ͺ]d)SIRx,SIAzQjcJˊ X73-J SxRyuo?.ވ#imҽkq+">N7ΕskNk gTK2ڛZ|sao8;| 3y,Cc87w\Q[T*d*k牔t6Bk)9*}mT^Z2؟uPdէIى%GɛrAΞo3a1DNvGҨeSDyYc.=VԔks'Q-[eԽD%o(vCzJJB 拉%DԒ6`0),H/1YxK Fy$*9m!uͬRD7arUI=;ҫ. :s~nR/}<ZzjUtf.\ea^>FÛpf}g N/r=UN.,ˢ4MAW̥-p~dIScn'[Qm8KKwHu$u㕦[I/򇰤v_*)PRߟxi'7ȅfZ n@ zzcsYy)y&^´vP`+)m! di(o|+uGf~F ^ ^tAHmh'>_X\0]u'PK!1drs/downrev.xmlDKk1FCB7faG F+bwɝN!8E8eP uItndKǯ%6F,9fzO/Yh6)tӒ0V-GjSr (:%vK؝/0=oލ㰚;>j+cK>+>'+ ܈/~PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!;F⺳ *drs/shapexml.xmlPK-!1 drs/downrev.xmlPKq2   j*   a L  r0e0e?Rectangle 2R   [ L  ~sy˼y˼?Rectangle 3 31  [ )For example, we can use existing tools to automatically verify: when is it safe to replace one system with another? We can automatically check whether models have the same possible sequence of interactions with their environment. [These three models are all bisimilar (weak open bisimulation) But if we cut one of the back edges they are no longer bisimilar.] So we have a way of splitting a large model into simpler parts, and analysing these parts individually.     33 ?   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The calculus is essentially defined in terms of actions and processes. Each process can be used to describe the behaviour of a component in the system, such as a gene or protein, while the actions describe what the component can do. (Assume a calculus with only bound output for the purposes of this talk) Delay represents an internal reaction such as radioactive decay, or change of shape. Each delay is associated with a rate, which determines the average duration of the reaction. Two components interact by performing a complementary send and receive on the same channel. Can represent two proteins with complementary shapes, or two chemicals that are known to react with each other. Note that 3-way reactions are extremely rare: very low probability of three molecules reacting at exactly the same time. Usually two molecules interact, and then a third. Pi-calculus fits this model nicely. Often a given molecule can do more than one thing. E.g. can either react with another molecule or decay. Represented as a choice of actions. We can of course have multiple molecules in parallel. Represented as parallel composition. We also want to define different species of molecules, using parameterised definitions in an environment. Finally, we want to create private bonds to represent the formation of complexes. A system S consists of a top-level environment E together with a process P.   `  0e0e?4Slide Image Placeholder 6R   [ 33h (  d(  d; d fCy˼y˼?Rectangle 7""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!R_$ drs/shapexml.xmlUMo6 x-l_XeqZ 8EϔDŪ)R%)No(9EQy(o޼~t(K c}q&ujB?eg™BgB-c~U k7bu.R{Xڧne Jzn) ͯJ7ڒזDdę%})P<)Ƽ:7nވ#ieҝkq+~isgUbsZ"s 9G_L&@<䶼j!a4 Wc>h2!tb&pD^*U/f<+4b( y"|ͲPR BJ_G@*A^ҡT^" 0;R(y_/>t#hٳСVr~֐t4A>, v'yraTв)<1+t|jJT驓߄ڲzlW7Qo={!%i"jRUHC0BgQ rQ$ŬoX Fy$*9m!uͬRD7arUN= ҫ72t 7|ޝݤ^(x&\"=~d1ͤ \Q¼sM{|Lnǝ`cM;?NWU?+C p]AKUeQߋ +RwnphdIScn'[QM8IKwHu$u㕦[I/򧰤v_()PRߟxi'7ȅfZ n@ zzCsYy)y"^´vP`+)m! di(o|~(uGf~F ^ ^׌tAHho'>Y<_0]uPK! L/drs/downrev.xmlDOk1GCRzۭQD-xJ۸C7ɒdo7o aw6Zt#YRksh0rL^~%2Pyf"nbZґ*%(W %ɜ+u\MEϩS谹owWunx0O?yT{[[:V솛'+ ܈/PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!R_$ *drs/shapexml.xmlPK-! L/ drs/downrev.xmlPKq2   j*   a d  ~Cy˼y˼?Rectangle 3 31  [ We developed a simple graphical representation for the calculus. Assume a single constraint on the syntax: each choice is defined as a species in the environment. A choice pi_1.P1 +. . .+ pi_N.PN is displayed as an elliptical node with directed edges to processes P1, . . . , PN. Each edge to a process Pi is labelled with an action pi_i and denotes an alternative execution path in the system. A parallel composition P1 | . . . | PN is displayed as a rectangular node with directed edges to processes P1, . . . , PN. Each edge to a process Pi denotes a concurrent execution path in the system. A species X(n) with definition X(m) = P is displayed as a rectangular node with a directed edge to the process defined as X(m). The tip of the edge is labelled with the substitution m:=n. Identical pairs in the substitution are not shown, while empty substitutions where n = m are omitted altogether. Restricted names are displayed by placing them next to the node of the process. Processes in the environment are displayed by connecting edges to nodes based on the node identifiers Xi(mi). 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Links are drawn from private channels to substitution tokens in order to represent the formation of complexes. Allows a graphical execution model. 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(5F(kKVv_[V2fcѢ¹2'%ل;G8V&۹7r+_ 1m,tlq\XkR^A?`=@ݾ<^Ɂ_4VTMQ0 xcg`8:128h*f&h1sBBuNQ1 sJL~TR BJ_Ut@@*A^ҡT^" 0?R(y_/>t#hٳСVr~Ґ aMc,۝nĨe3DyYk&+XfZ٩_zlW7Qo={!%i"jJUHC0RQ rQŬoXj "| Gœ6Y7:CvV)0:[gUUAhg, ߟwn7meִټHuqߩ?m;YC;tT0,g$Y̖fNzqrӛ-&hEXc:U ZʮV_1wc%K*gs t;rlNZp@?#4J|?%Lt5|@A@I}_ज़ ix%meKf; ABBd@PHh+A8R=)w?&zXO{_3ZO4 uVmi4~f=Ԣ|UtPK!,drs/downrev.xmlDMK1@!El*ҮMK)+ZDon%vgX%ӊ 8a [8J}{?"Z`s{1V%6䨠˥ EMud9|Ͼ p.cPK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!l *drs/shapexml.xmlPK-!, drs/downrev.xmlPKq2   j*   a |  ~@y˼y˼?Rectangle 3 31  [ Two molecules can bind by sending and receiving on the same channel, where the argument represents a new channel that can be used to split the complex.    |  0e0e?4Slide Image Placeholder 6R   [ 33    z (     0e0e?4Slide Image Placeholder 1R   [    y˼y˼?(Notes Placeholder 2 31  [ 6  X  y˼y˼?6Slide Number Placeholder 3"!PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!ot drs/shapexml.xmlUQo6~@pmَcU ۫^`LIT"5r 4.<(GxJҺGo{I1~sЙPF˘o~]5uaU̷Wnץ[Y TRc/7KЭtR{qQ^o-E BZ[һڲ"ję%ݨ".iZTn`x=G!?Vʤ;_3iskdDWu\ ͏H7pڛR sp0IE1Sy, ^ dؿ ŔgebTԇL~_CVE^ҡT^R)-;RQ^eBи5G+ 8w-L<"|X ǃNrG(-:EԘ''sB_MZjySت-f+*_m|CPӔjb d)* c0BgMsr^$5YxKs FY8G|6o~4Y:Ef()0*]UYAhg_ ϻ_ e֤<=Kào՟4,`c3tU0h0wɰ3f|G|K4^~X +CGs Zʮ.U4_1s;dzڒ%M·@zlCDm6l'-axZ m֑ԍ4Fx%L5&+BIP4K; A.4I╗sLJ[9^l! di(/|ޔ#E3Rj#uG/ ߴ #]EcRgkaƉ3n>PK!@kcdrs/downrev.xmlDAK@Fa -EkA0m.4IΆm{z>|9N |kY4A\YrܽLAL ac Q>GM}.2觶'loOWK妓$ɤB==5T}oF].=>l0J]_G>?Z2K Z]1PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!ot *drs/shapexml.xmlPK-!@kcdrs/downrev.xmlPKq2   j*   a 33 umU(  ;  f`y˼y˼?Rectangle 7""PK!Zf[Content_Types].xmlMO &2WR=cJ`F0iK`#̼vLw 9uSq:w`G ^i ½KI)c/ $oVjTMRc|}042ҥCƔM̏P~*ka/8^DkHbL8e i"K\XN\6rco4y@_;oPK!1_a _rels/.relsj0 ѽqCNo^K [ILcX&m߾0XFo;>0xM e`|X}đ I`߽N4aG2$RKIZ)4(M9`ctB{m:f@`3n|O,ܗr޾jxR0T ,0@}WBLǬ5vPK!  drs/shapexml.xmlUn6?^[c* mSLKT"Ur8 X=P ߼y3P)֕F<0L~JϏ1g eLQ:o>SW3|ݴNzl++>ZjVciNj/<Tj0+Qj~PzW~́%'gZT8Af@$Kxso?ވ#iip+"ʙy+E5l s*m29ƛPR|) 0kΎ)DWh<&tb*epdpY<_0]}'PK! L/drs/downrev.xmlDOk1GCRzۭQD-xJ۸C7ɒdo7o aw6Zt#YRksh0rL^~%2Pyf"nbZґ*%(W %ɜ+u\MEϩS谹owWunx0O?yT{[[:V솛'+ ܈/PK-!Zf[Content_Types].xmlPK-!1_a /_rels/.relsPK-!  *drs/shapexml.xmlPK-! L/ drs/downrev.xmlPKq2   j*   a   ~ y˼y˼?Rectangle 3 31  [ >We developed a simple graphical representation for the calculus. Assume a single constraint on the syntax: each choice is defined as a species in the environment. A choice pi_1.P1 +. . .+ pi_N.PN is displayed as an elliptical node with directed edges to processes P1, . . . , PN. Each edge to a process Pi is labelled with an action pi_i and denotes an alternative execution path in the system. A parallel composition P1 | . . . | PN is displayed as a rectangular node with directed edges to processes P1, . . . , PN. Each edge to a process Pi denotes a concurrent execution path in the system. A species X(n) with definition X(m) = P is displayed as a rectangular node with a directed edge to the process defined as X(m). The tip of the edge is labelled with the substitution m:=n. Identical pairs in the substitution are not shown, while empty substitutions where n = m are omitted altogether. Restricted names are displayed by placing them next to the node of the process. Processes in the environment are displayed by connecting edges to nodes based on the node identifiers Xi(mi). The notation Pi*Xi(mi) denotes a process Pi with label Xi(mi), where the label is placed inside the node of process Pi A running process is displayed by placing a substitution token next to the relevant nodes in the graph. Links are drawn from private channels to substitution tokens in order to represent the formation of complexes. Allows a graphical execution model. 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   ( On-screen Show (4:3)Microsoft CorporationH{pn zArial WingdingsTimes New RomanSymbol Courier New MS PGothic MS MinchoTEMPLATE_Faculty_Summit_20061_TEMPLATE_Faculty_Summit_2006Package7A Visual Programming Language for Biological ProcessesBiological ComputingSystems Biology#Large, Complex, Biological Models Biological Programming "Programming Languages for BiologyLanguage DevelopmentSPiM: Stochastic p for BiologyMessage-Passing ApproachCompact, Modular Models"EGFR Model [Hornberg et. al 2005]"EGFR Model [Hornberg et. al 2005] Modular EGFROutlineBasic SPiM Examples4Production: G produce G + P P degrade 4Production: G produce G + P P degrade 4Production: G produce G + P P degrade 4Production: G produce G + P P degrade 4Production: G produce G + P P degrade 4Production: G produce G + P P degrade Gene Simulation#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + YpBinding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Programming Gene Networks*Repressilator [Elowitz and Leibler, 2000]Parameterised Gene Gate*Repressilator [Elowitz and Leibler, 2000]Graphical ProgrammingHow does the oscillator work?Oscillator: 0sOscillator: 5.568177sOscillator: 6.329912sOscillator: 11.62149sOscillator: 13.21617s Oscillator Oscillator OscillatorOscillator: 11039.31sOscillator: 11039.77sOscillator SimulationAnalysing Simulation TracesRepressilator TraceA Gene Gate in 3DThe Repressilator in 3DGraphical DebuggingParameter AnalysisModel Refinement:"Cooperativity improves robustnessCooperativity Simulations)Bacteria Logic Gates [Guet et al., 2002]Gene with protein inhibitorBacteria Logic Gates7A Computational Model of C. elegans Vulval DevelopmentC. elegans Nematode WormC. elegans VPC DifferentiationSimplified Model of AC and VPCNetwork of AC and 6 VPCPopulation PlotGeometric PlotRefined VPC ModelSimulation Results:A Computational Model of MHC class I Antigen Presentation MHC: A Biological Virus Scanner MHC: A Biological Virus Scanner Investigate the Role of TapasinMHC I StructureSPiM Peptide Editing ModelModel ParametersSimulations Match ExperimentsPeptide DiscriminationParameter SpacePeptide DiscriminationParameter SpacePeptide Loading: Flytrap ModelFlytrap Peptide Editing Model Peptide Discrimination: FlytrapKey mechanisms identifiedMHC Alleles: Model PredictionsExtending the ModelExtendable,Maintainable ModelsVerifying Biological ModelsModelling Immunodominance References ReferencesSPiM Definition SPiM Syntax Graphical Syntax: Environment EGraphical Syntax: Process PGraphical Semantics: DelayGraphical Semantics: Delay!Graphical Semantics: Interaction!Graphical Semantics: InteractionGraphical Semantics: BindingGraphical Semantics: BindingGraphical Syntax  Fonts UsedThemeEmbedded OLE Servers Slide Titlesp _="/aphillipaphillip>>>>>>>>>>>>>>>>>  !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      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PowerPoint Document(,="DocumentSummaryInformation8>MsoDataStore ǵFǵFHCXQ3VEMGSQ==2ǵFǵFItem  PropertiesItem >Properties ՜.+,D՜.+,D    ( On-screen Show (4:3)Microsoft CorporationH{pn zArial WingdingsTimes New RomanSymbol Courier New MS PGothic MS MinchoTEMPLATE_Faculty_Summit_20061_TEMPLATE_Faculty_Summit_2006Package7A Visual Programming Language for Biological ProcessesBiological ComputingSystems Biology#Large, Complex, Biological Models Biological Programming "Programming Languages for BiologyLanguage DevelopmentSPiM: Stochastic p for BiologyMessage-Passing ApproachCompact, Modular Models"EGFR Model [Hornberg et. al 2005]"EGFR Model [Hornberg et. al 2005] Modular EGFROutlineBasic SPiM Examples4Production: G produce G + P P degrade 4Production: G produce G + P P degrade 4Production: G produce G + P P degrade 4Production: G produce G + P P degrade 4Production: G produce G + P P degrade 4Production: G produce G + P P degrade Gene Simulation#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + Yp#Interaction: Xp + Y da X + YpBinding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Binding: X + Y -b+b X'Y'Programming Gene Networks*Repressilator [Elowitz and Leibler, 2000]Parameterised Gene Gate*Repressilator [Elowitz and Leibler, 2000]Graphical ProgrammingHow does the oscillator work?Oscillator: 0sOscillator: 5.568177sOscillator: 6.329912sOscillator: 11.62149sOscillator: 13.21617s Oscillator Oscillator OscillatorOscillator: 11039.31sOscillator: 11039.77sOscillator SimulationAnalysing Simulation TracesRepressilator TraceA Gene Gate in 3DThe Repressilator in 3DGraphical DebuggingParameter AnalysisModel Refinement:"Cooperativity improves robustnessCooperativity Simulations)Bacteria Logic Gates [Guet et al., 2002]Gene with protein inhibitorBacteria Logic Gates7A Computational Model of C. elegans Vulval DevelopmentC. elegans Nematode WormC. elegans VPC DifferentiationSimplified Model of AC and VPCNetwork of AC and 6 VPCPopulation PlotGeometric PlotRefined VPC ModelSimulation Results:A Computational Model of MHC class I Antigen Presentation MHC: A Biological Virus Scanner MHC: A Biological Virus Scanner Investigate the Role of TapasinMHC I StructureSPiM Peptide Editing ModelModel ParametersSimulations Match ExperimentsPeptide DiscriminationParameter SpacePeptide DiscriminationParameter SpacePeptide Loading: Flytrap ModelFlytrap Peptide Editing Model Peptide Discrimination: FlytrapKey mechanisms identifiedMHC Alleles: Model PredictionsExtending the ModelExtendable,Maintainable ModelsVerifying Biological ModelsModelling Immunodominance References ReferencesSPiM Definition SPiM Syntax Graphical Syntax: Environment EGraphical Syntax: Process PGraphical Semantics: DelayGraphical Semantics: Delay!Graphical Semantics: Interaction!Graphical Semantics: InteractionGraphical Semantics: BindingGraphical Semantics: BindingGraphical Syntax  Fonts UsedThemeEmbedded OLE Servers Slide Titlesp0PublishingExpirationDatePublishingStartDate 2019-03-12T12:09:07Z2000-01-01T00:00:00Z DocumentLibraryFormDocumentLibraryFormDocumentLibraryForm This value indicates the number of saves or revisions. 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