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Pb$8oE"\"$^j߅U(z"$d.] Rܼ ~}>ң2$(" %W*gO2$[d[Kݣr* 0AA@ʚ;(74ʚ;g4zdzd< 0ppp@ <4dddd@w 0t0 0___PPT10 pp~___PPT9`?  %,:Protein traffic control Mikael Bodn mikael@itee.uq.edu.auT; && hAbstract The cell is a decentralised but still carefully controlled device, shuttling gene products, like proteins, through various compartments where they perform their functions. Mechanisms for this protein traffic control are not yet fully understood and machine learning techniques are being utilised to discover protein sequence signals that allow targeted drug design and the development of models that can be used to automatically annotate the growing number of sequences that are yet to be experimentally characterised. In relation to the current state-of-the-art, we report on improvements in accuracy for a range of models for predicting localisation of proteins -- making use of advanced machine learning techniques like recurrent neural networks and support vector machines. In particular we focus on the essential compartments and organelles, the secretory pathway, the mitochondrion, the chloroplast and the peroxisome. Our models are housed in the Protein Prowler, an online sequence prediction server, which has been endowed with graphical features allowing the user to scrutinize the outputs of the models. 0 P`P `Air traffic control Modelling problemhATC: the system is engineered, models are used to simulate/improve the design further PTC: the system is a result of evolution, models are used to understand the existing system A cell can contain 10,000 different kinds of proteins The eukaryotic cell has evolved several functionally specialised compartments and membranes, including a pathway for protein secretion/export, a membrane-lined nucleus, mitochondria, peroxisomes, & But most proteins are synthesised on ribosomes in the cytosol& how do they transit? Let s look closer at the main principles& [movie] PWPPP W!  "9 0-2 Protein traffic controlFAre there sequence zip-codes? Well& $$ Signal peptides (SP) Target proteins to the secretory pathway (the ER initially). 15-30 residues long. N-terminal domain positively charged, central hydrophobic region, small apolar residues -3, -1 relative cleavage site. Mitochondrial targeting peptides (mTP) Target proteins to mitochondria (matrix, outer and/or inner membrane) 20-40 residues long. Arg, Ala and Ser overrepresented, while negative residues scarce. Weak conservation around cleavage site(s). Chloroplast transit peptides (cTP) Target proteins to chloroplast. 20-120 residues long. Hydroxylated residues common, acidic uncommon. Semi-conserved motif around cleavage site. Peroxisomal targeting sequences (PTS) Target proteins to the peroxisome Either a short C-terminal pattern or a vaguely defined N-terminal pattern. PP'PP#PP&PmPP' <]# r&       K    b!8 N  VPredicting localisation from protein sequence data: Some methodological considerationsWW7Expert systems (utilising known principles) are clearly outperformed by many machine learning models trained on sequence data. Sequence data: the sequence of amino acids that make up the protein variable length several hundreds amino acids long (consider 100 a.a. long=20100 combinations!) can be analysed by amino acid composition a pre-specified fixed part of the sequence a sliding window (a stretch of amino acids) where each a.a. is encoded numerically (usually a vector) Several hundreds, if not thousands of sequences, with experimentally verified localisation, constitute training and test data. Training and test data are further checked so that redundancy (prevalent in data banks) is kept to a minimum. Models and algorithms Feed forward neural network Bidirectional recurrent network Support vector machine >PPDP2PAPnP;PPPPSPPDM n:      S, #Detect N-terminal targeting signals$$(HSignal peptides, mitochondrial targeting peptides, chloroplast transit peptides, are located in the N-terminal end of the sequence (and are often cleaved/removed after transit). Sliding window approach works well, producing scores of  signal presence . TargetP is the most reliable predictor. #Detect N-terminal targeting signals$$( #Detect N-terminal targeting signals$$(~Recurrent networks demonstrate ability to make accessible, and find sequential targeting signals (overall improvement of ~20% on non-plant data, ~15% on plant data, compared to TargetP) Differences in ability of architectures prompt development of predictors utilising combinations. A combined target classifier is 6% better on non-plant data, and 5% better on plant-data, compared to TargetP (as measured by Matthews correlation coefficient). +ZZZ+A   Y #Detect C-terminal targeting signals$$(yPeroxisomal localisation uses mainly a C-terminal targeting signal (PTS1) PTS1 is often described as a conserved 3-a.a. sequence (usually Serine-Lysine-Leucine) at the C-terminus. However, recent studies show dependencies to preceding a.a. A curated dataset was extracted from SWISSProt (positives and negatives, redundancy-reduced, and erroneously annotated sequences removed)zZzH  \#Detect C-terminal targeting signals$$(#Detect C-terminal targeting signals$$(#Detect C-terminal targeting signals$$(New larger dataset Exploiting the full range of dependencies at the C-terminus Overall ~25% better classification accuracy and higher specificity (0.91 vs. 0.64) than previously reported studies. @OvO j3The Protein Prowler (http://pprowler.imb.uq.edu.au)44(" 03xA prediction service for determining the localisation of proteins from sequence data. Currently, the software distinguishes between Secretory pathway Mitochondria Chloroplasts Other Peroxisome and nucleus will be integrated. Handling of membrane proteins. Backend algorithms developed, evaluated and reported in the literature (see references). Graphical features were developed (supported by ACCS)  allowing visual inspection of some results.vVZ.Z2ZJZZV.2J   Where next?So far modelling of targeting processes, system-level modelling would capture dynamics of shuttling, e.g. nuclear export of proteins only happen once they have been imported. More comprehensive data sets, including membrane proteins (notoriously difficult, but very relevant).6FifFifAcknowledgmentscJohn Hawkins Stefan Maetschke Mark Wakabayashi James Watson Rohan Teasdale Zheng Yuan Melissa DavisdZd, . ITEE and ACCS IMB and ACBZDReferences Download from http://www.itee.uq.edu.au/~mikael/publ.html2E ( (,"$ 0D/p   0` 33` Sf3f` 33g` f` www3PP` ZXdbmo` \ғ3y`Ӣ` 3f3ff` 3f3FKf` hk]wwwfܹ` ff>>\`Y{ff` R>&- {p_/̴>?" dd@,|?" dd@   " @ ` n?" dd@   @@``PR    @ ` ` p>> $(    6䑒  `}  T Click to edit Master title style! !  0Д  `  RClick to edit Master text styles Second level Third level Fourth level Fifth level!     S  0 ^ `  X*  0Ƞ ^   Z*  0 ^ `  Z*H  0޽h ? 3380___PPT10.R@3 Default Design 0 zr@ (    0, 86   P*    0 o 6  R*  d  c $ ?Dd    0X  n  RClick to edit Master text styles Second level Third level Fourth level Fifth level!     S  6  8?   P*    6  o ?  R*  H  0bgy ? 3380___PPT10.R@zO$ 0  $(  r  S 4p 4 r  S 4`9 4 H  0޽h ? 3380___PPT10.R,3<  0 p <(   r  S Y9 `}  9   C bAHair-traffic-control-baytracon-west2zRH  0޽h ? 3380___PPT10.UpG  0 (*(  (r ( S ]9 `}  9 x ( c $^9 ` 9 H ( 0޽h ? 33___PPT10i.Um+D=' 4= @B +[  0 rj,(  ,r , S e9 `}  9 P , C (Ach17f1zt `H , 0޽h ? 33___PPT10i.U@.+D=' 4= @B +  0 0 (   x  c $:9F   9 x  c $:9Dm < 9   <U d 2From Emanuelsson, Brief. Bioinformatics 3(4) 2002.*3  #X  0A?p H  0޽h ? 33___PPT10i.+D=' 4= @B +$  0 8$(  8r 8 S lp9I'g  9 r 8 S q9 ` 9 H 8 0޽h ? 3380___PPT10.Uu$  0 p$(  pr p S 9 `}  9 r p S l9 ` 9 H p 0޽h ? 3380___PPT10.UPh'  0  <(  <2F D< <  qi < 0C"?D<^ < c .A bifnS   < <v9"`u I 0 l& VRACNQA& > ^F D< < D< < 0C"?D< < <A ??  x  < c $9 `}  9   < <9 pT  .Targeting signal detection  < <9p 1Overall target classificationH < 0޽h ? 33___PPT10i.9+D=' 4= @B +0  0 l0(  lx l c $h9 `}  9 x l c $@9 ` 9 H l 0޽h ? 3380___PPT10.U^^$  0 h$(  hr h S Ķ9 `}  9 r h S 9 ` 9 H h 0޽h ? 3380___PPT10.UpCb  0 xb(  xZ x C 2Adependencies\r x S x9 `}  9  x0 NA  ? ?l 7N   9RB x s *D z  RB x s *DF z @ H x 0޽h ? 3380___PPT10.U@M   0 n f (  r  S 9 `}  9   <P30044 MGLAGVCALRRSAGYILVGGAGGQSAAAAARRCSEGEWASGGVRSFSRAAAAMAPIKVGD AIPAVEVFEGEPGNKVNLAELFKGKKGVLFGVPGAFTPGCSKTHLPGFVEQAEALKAKGV QVVACLSVNDAFVTGEWGRAHKAEGKVRLLADPTGAFGKETDLLLDDSLVSIFGNRRLKR FSMVVQDGIVKALNVEPDGTGLTCSLAPNIISQL $    0 9~  : 2   <<9"`}Id zCheck for SP, mTP, cTP,RB  s *D  }  0@9"` z  3SVM  6x9"`P $  B TCSLAPNIISQL  RB  s *D`  CRB  s *D%      <9"` ] j  Yes  09"` j  2NoRB  s *DFF RB  s *D   LB  c $DCFLB  c $Dc Fc H  0޽h ? 3380___PPT10.U ]h$  0 $(  r  S e `}   r  S f `  H  0޽h ? 3380___PPT10.U}  0 H$(  Hr H S l `}   r H S m `  H H 0޽h ? 33___PPT10i.Uؓ+D=' 4= @B +$  0 @$(  r  S   `}   r  S < `  H  0޽h ? 3380___PPT10.tV?!w  0 `(  r  S <4 `}  4 r  S 4  6  4 r  S 4p ` 4 H  0޽h ? 33___PPT10i.R0zs+D=' 4= @B +  0  '(  r  S ȇ `}    ' 6] } Bodn, M. and Hawkins, J. Prediction of subcellular localisation using sequence-biased recurrent networks. Bioinformatics. In press. Wakabayashi, M., Hawkins, J., Maetschke, S. and Bodn, M., Exploiting sequence dependencies in the prediction of peroxisomal proteins. In Proceedings of IDEAL 2005, Brisbane. In press. Hawkins, J. and Bodn, M. The Applicability of Recurrent Neural Networks for Biological Sequence Analysis. IEEE/ACM Transactions on Computational Biology and Bioinformatics. 2(2), 2005. Bodn, M. and Hawkins, J. Improved access to sequential motifs: A note on the architectural bias of recurrent networks. IEEE Transactions on Neural Networks. 16(2), 491-494, 2005. Bodn, M. and Hawkins, J. Detecting residues in targeting peptides. In Proceedings of the Asia-Pacific Bioinformatics Conference. Singapore. 2005. Maetschke, S., Towsey, M. and Bodn, M. BLOMAP: An encoding of amino acids which improves signal peptide cleavage prediction. In Proceedings of the Asia-Pacific Bioinformatics Conference. Singapore. 2005.  2kA$_99  < M  H  0޽h ? 3380___PPT10.U~ 0 LDP(  d  c $Dd   8  s * n   Sorting relies on signal/targeting sequences. Often short sequences located at the N-terminus (signalling  export through secretory pathway,  import into mitochondria and chloroplast). Peroxisome and nucleus import different (folded, not N-terminal). Here, schematic characterisations of targeting sequences grouped according to destination.H  0bgy ? 3380___PPT10. 0 `X@(  @d @ c $Dd   L @ s * n   tHow does TargetP operate? Similar to SignalP, which scans/slides over the sequence, taking snapshots, and produces an  s-score ( signal peptide-ness ) for each residue. The s-scores are used collectively to say whether the sequence has a signal peptide. Distinguishes only between sequences with a signal peptide and those without. TargetP comes in two versions: one plant and one one-plant. Both employ target-specific detection networks scanning the sequence and then A sorting network which makes the sequence classification. Window sizes? Training? Testing? (}#;&H @ 0bgy ? 3380___PPT10.T. 0 D>(  D^ D S Dd    D c $ n   4Hormone receptor proteins, for example, must be delivered to the plasma membrane if the cell is to recognize hormones, and specific ion-channel and transporter proteins are needed if the cell is to import or export the corresponding ions and small molecules. Water-soluble enzymes such as RNA and DNA polymerases must be targeted to the nucleus; still others, such as proteolytic enzymes or catalase, must go to lysosomes or peroxisomes, respectively. (Lodish et al, ch 17).> &H D 0bgy ? 3380___PPT10.Up 0 kc L(  L^ L S Dd   ] L c $ n   P27456, Glutathione reductase, and P29463, CoxVa/Triose phosphate translocator), and the outputs of two individual networks. P27456, more frequently found in the chloroplast (only 3% in mitochondria), has a putative cleavage site at position 60, which is well-recognised by the recurrent cTP detector. 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Mechanisms for this protein .-@Arial-. 2 4traffic .-@Arial-. g2 8@control are not yet fully understood and machine learning techni.-@Arial-. 2 8ques i.-@Arial-. g2 ;@are being utilised to discover protein sequence signals that all.-@Arial-.  2 ;|ow .-@Arial-. f2 >?targeted drug design and the development of models that can be l.-@Arial-. f2 B?used to automatically annotate the growing number of sequences l.-@Arial-. Q2 E1that are yet to be experimentally characterised. s.-@Arial-. 72 N In relation to the current state.-@Arial-.  2 NI-.-@Arial-.  2 NJof.-@Arial-.  2 NM-.-@Arial-.  2 NNthe.-@Arial-.  2 NS-.-@Arial-. 62 NTart, we report on improvements .-@Arial-. g2 Q@in accuracy for a range of models for predicting localisation of.-@Arial-. 2 U proteins .-@Arial-.  2 U(--.-@Arial-. [2 U+8making use of advanced machine learning techniques like .-@Arial-. g2 X@recurrent neural networks and support vector machines. In partic.-@Arial-. 2 Xular i.-@Arial-. `2 [;we focus on the essential compartments and organelles, the a.-@Arial-. d2 _>secretory pathway, the mitochondrion, the chloroplast and the .-@Arial-. g2 b@peroxisome. Our models are housed in the Protein Prowler, an onl.-@Arial-.  2 bine .-@Arial-. g2 f@sequence prediction server, which has been endowed with graphica.-@Arial-.  2 fl .-@Arial-. g2 i@features allowing the user to scrutinize the outputs of the mode.-@Arial-.  2 i~ls. .-՜.+,D՜.+,0    eOn-screen ShowITEE+G ArialTimes New Roman Courier NewCourierDefault DesignMicrosoft Visio DrawingMicrosoft Equation 3.0;Protein traffic control Mikael Bodn mikael@itee.uq.edu.auAir traffic controlModelling problemProtein traffic control$Are there sequence zip-codes? WellWPredicting localisation from protein sequence data: Some methodological considerations$Detect N-terminal targeting signals$Detect N-terminal targeting signals$Detect N-terminal targeting signals$Detect C-terminal targeting signals$Detect C-terminal targeting signals$Detect C-terminal targeting signals$Detect C-terminal targeting signals4The Protein Prowler (http://pprowler.imb.uq.edu.au) Where next?AcknowledgmentsEReferences Download from http://www.itee.uq.edu.au/~mikael/publ.html  Fonts UsedDesign TemplateEmbedded OLE Servers Slide Titles 8@ _PID_HLINKSA7C:\Documents and Settings\mikael\Desktop\ch17anim1.movhttp://pprowler.imb.uq.edu.au/,http://www.itee.uq.edu.au/~mikael/publ.html_㻒0ITEEITEE  !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~Root EntrydO)PicturesCurrent UserSummaryInformation($PowerPoint Document(ߒDocumentSummaryInformation8