ࡱ>  |)`0bjbjWwWw  55I; >>>R:DRVTTT999UUUUUUU$Wh%ZbU>BQ809BBUTVJJJBT^>UJBUJJVjR@> S~ `dGR UV0VRTZ7IZ S SZ>S9_< Jk>@999UUJ999VBBBBRRRDRRRRRR }elimir Kurtanjek, Hrvoje }ili, Ana Jurinjak Tuaek University of Zagreb, Faculty of Food Technology and Biotechnology Pierottijeva 6, 10000 Zagreb, Croatia Systems biology and biotechnology ABSTRACT Systems biology in present genome era receives the most important position in modern genome based biotechnology. Its importance is result of exceptional development of experimental high-throughput techniques which deliver high dimensional multivariate data in a single biological experiment. Availability of large data sets are sources of biologically essential information on a system level which can be only inferred by mathematical systems theory concepts and application of mathematical/statistical methods and bioinformatics tools. The integration of information derived from a genome of industrially important microorganisms and bioprocess engineering has led to the new generation of genome based biotechnology. Applications are very diverse, from the fundamental research in biology to applicative fields such as medicine, ecology, and most importantly in industrial biotechnology. Here is given a short review of the basic concept of the biological systems theory from a view point of biochemical engineering with the aims of white (energy production) and red (pharmaceutical production). Case studies are presented results of the system flux control analysis of E. coli glycolysis and the S. cerevisiae pathways for ergosterol and sphingolipid synthesis. Key words: systems analysis, metabolic networks, global sensitivity, CellDesignerTM INTRODUCTION Mathematical theories of biological systems have developed in parallel with experimental biology but initially had relatively limited impact on the science of biology. However, with the start of genome era and especially the development of many high-throughput techniques with parallel and robot assisted systems, mathematical systems theories based on experimental molecular biology become highly developed. Most of the biological data are shared on internet in form of various open source data basis and are constantly being updated and corrected by the global scientific community. With rapid advancement of experimental techniques based on molecular tools, robotics and informatics, numerous industrially important microorganisms (about 200) have genome based (bottom-up) mathematical models of reconstructed metabolic networks.1 In order to extract information and theories from the vast quantity of data developed are mathematical and statistical tools (chemometrics and bioinformatics) for statistical inference and data structure analysis. Simultaneously are in development mathematical theories of living systems, which in the field of biotechnology are mostly models of industrially important microorganisms as producers of energy and bulk chemicals (white biotechnology) and pharmaceuticals (red biotechnology). This biology system model approach has led to the new paradigm of genome based biotechnology to emphasize full potential of microorganism biological potential and a systems approach to genome engineering. The main concept is schematically presented in Fig. 1 in which a model of a producing microorganism can be approached from the top to down or the macro (organism) system level, or from the bottom to up or the molecular level. Integration and synergism of the two approaches may be the most rational road map. Although the whole field is developing at exponential rate, a fully functional and validated living cell model (living cell in silico) on a computer is not yet available. For most of industrially important microorganisms (about 200) have available genomes which form the basis for development of their corresponding reconstructed metabolic networks. The knowledge of the reconstructed metabolic networks together with thermodynamic balance of free Gibbs energy for each of biotransformation reactions leads to determination of a space of feasible phenotypes.  SHAPE \* MERGEFORMAT  Figure 1. Schematic representation of the concept of genome based biotechnology mathematical model paradigm. N represents stoichiometric matrix of a reconstructed metabolic network, C are intracellular concentrations, F are cell input and output fluxes, and J are life goal (cybernetic) controlled intracellular fluxes (reaction rates). A genome based metabolic reconstructed metabolic network are inferred from whole annotated genome, biological information and thermodynamic constraints is stored in a large metabolic stochiometric matrix N. Application of linear algebra in analysis of N provides detailed structural properties of a network topology and biological potential. Intensity of flows of energy and mass through the network are determined by thermodynamic laws but the flows are controlled based on a signaling system based on the top to down approach derived from the basic cybernetic principles for living systems. These two approaches are integrated into a computer model in a form of a large set of intrinsically nonlinear differential equations (Fig. 1) which represent mass balances of intracellular components connected to environment through a cell trans-membrane fluxes. Detailed analysis of these systems based models reveals new emerging properties which together with genetic engineering enables rational strategy for development of a new generation of industrial biotechnology. SYSTEMS VIEW CELL MODEL Impacts of computer models and bioinformatics have dominant positions on genome based biotechnology. The central roles are genome based metabolic network reconstruction, enzyme kinetic models and multilevel regulatory mechanisms. Dynamical transient response of intracellular metabolite are given by:  EMBED Equation.3  (1) where N is a stochiometric matrix of a genome level reconstructed network, J are intracellular fluxes, F are trans-membrane fluxes of molecule exchange between a cell and its environment, and c and E are metabolite and enzyme concentrations. Topological properties of the network are based on linear algebra algorithms for evaluation of potential capabilities of a specific cell. Even more importantly, is the possibility to test in silico various redesign combination of a specific cellular mechanism by genetic engineering manipulation. Overall specific metabolism intensity is expressed by cell specific growth rate EMBED Equation.3 :  EMBED Equation.3  (2) The mass balances eq. 1 are tied up to hierarchical goal driven, but very intricate and complex, mechanism of genome transcription and regulation mechanism which could be possibly mathematically modeled on basis of systems cybernetic concepts (Fig. 1). Formally, it can be expressed by a biochemical reaction model based on the mass action principle mediated by cellular signaling network:  EMBED Equation.3  (3) The full genome scale models of the kinetic functions  EMBED Equation.3  and  EMBED Equation.3  are, at present experimental level far from reach and only fractional models for specific cellular functions have been proposed. From biochemical engineering standpoint, experimentally and theoretically possible are systems analysis based on models and data from homeostasis (chemostat) condition experiments and the corresponding FBA flux balance analysis. These models evaluate constant fluxes at different steady experimental conditions and importantly they do not involve error prone kinetic model functions and parameters. Homeostasis systems model has algebraic structure:  EMBED Equation.3  (4) Properties of the model equations, eq. 4, can be analytically fully analyzed and numerically solved, and for validated reconstructed networks are essential for genome based biotechnology. SYSTEMS BIOLOGY FOR WHITE AND RED BIOTECHNOLOGY Application of systems biology for industrial application is focused on a selection and definition of a basic chemical cell factory. Two main candidates are yeast S. cerevisiae and E. coli. Both microorganisms are in detailed studied and applied for various productions, and mathematical models of their reconstructed networks are available. Recently, when processes of white biotechnology are considered most of research available in literature is in favor of yeast primarily due to its robustness to harsh conditions in large volume bioreactors and less environmental and health risks. When the processes of a red biotechnology are concerned, E. coli due to its high metabolic activity provides higher productivities of low concentration metabolites in small bioreactor volumes with strict process control. Globally, the most important perspective of white biotechnology is production of a second generation of biofuels, i.e. production of fuels from lower grade bio-sources (agricultural waste rather then presently used cane sugar bagasse and corn), with higher energy content per unit volume, lower emissions, better miscibility with present fuels (gasoline), lower affinity to water and corrosion (J. Nielsen et al., 2013). The potentially main advanced biofuels are: ! butanols (four mixture with gasoline) ! sesquiterpentines (for jet fuel, diesel) ! fatty acid derived biofuels Presently is known that productivity of butanols in genetically modified S. cerevisiae exceeds that obtained in E. coli. Most of present industrial factories for production of bioethanol from corn due to the market changes resulting from surplus of hydrocarbons from shale gas will have to be retrofitted to low grade agricultural (cellulose) feed stock. E. coli has been proven as a cell factory for production of various recombinant proteins. Here is presented systems analysis results of E. coli central metabolism with Entner-Doudoroff shunt pathway (Tuek and Kurtanjek 2009; Kurtanjek 2010,). Analysis of the central metabolism is motivated by the fact that it is central control point for distribution of mass and energy throughout the whole metabolic network. It is also the most complex control point from which anabolic pathways spread out mostly as linear sequences of step by step transformations. Applied is the stochastic analysis of the metabolism control upon published data on glucose impulse. Control activity of each enzyme is evaluated under unsteady conditions during the first 10-15 seconds after imposed impulse under glucose deprived E. coli cell population in a fed-batch reactor5. The metabolic fluxes J are considered as a random variables defined by presumed probability distribution functions  for each enzyme E. Probabilistic global parametric sensitivity  (5) Flux control coefficients are evaluated as the global sensitivities determined by the conditional variances of the whole assembly of enzymes included into the model.  (6)  SHAPE \* MERGEFORMAT  Figure 2. Dynamic responses of the control flux coefficients for PTS mechanism and pyruvate to biomass flux. The computer simulations reveal the known fact that PFK enzyme is the control hub (focus) which undertakes the control from PTS enzyme complex in the very few fractions of a second. Further intake of glucose is solely controlled by PFK. However, fluxes around pyruvate and acetyl-coenzyme-A exhibit complex concerted control of multiple enzymes. These fluxes have distributed control and their redirection or amplification requires complex systems analysis.  SHAPE \* MERGEFORMAT  Figure 3. CellDesigner model of ergosterol and sphingolipds in S. cerevisiae (H. }ili, 2013). The second example given here is the study of interaction between pathways for synthesis of ergosterol and sphingolipds in S. cerevisiae (H. }ili, 2013). The model data (Alvarez-Vasquez, 2011) are given in the S-system form. The algebraic model equations are transformed to a CellDesigner model presented in Fig. 3 and also in Wolfram Mathematica software. Applied are dynamic local and global sensitivity analysis (by extended FAST method). Due to numerous fluxes (80) and parameters (147) the results are presented in a colored coded maps commonly called heat maps which are also frequently used for presentation of high-throughput experimental data. One of the obtained computer simulation scenarios is presented in Fig 4. The results indicate that there are 10 key controlling enzymes (of 80 included) which are in the initial parts of the synthesis pathways. Relatively small number of the controlling enzymes indicates the robustness of the synthesis.  SHAPE \* MERGEFORMAT  Figure 4. Heat map of dynamic flux control coefficients in the pathways for ergosterol and sphingolipds in S. cerevisiae. CONCLUSIONS From biochemical engineers perspective systems biology provides methodology and tools for systemic analysis of vast data available from biological experiments based on high-throughput experiments. Importance of the systems approach is ability to extract statistically significant information and relate the biological effects to external factors tested in a controlled and designed experiments. Systems level dynamic models provide computer simulations needed for optimization of existing technologies or design and optimization of new technologies in white and red biotechnology. Probably the most important results of biological systems models is their use for rational planning of genetic engineering simultaneous interventions of several key enzymes and possibility for development of synthetic enzymes tailored according to computer predicted objectives. LITERATURE 1. Ines Thiele, Bernhard . Palsson, A protocol for generating a high-quality genome scale metabolic reconstruction, Nat. Protoc. 5(1) 93-121(2009). 2. Jens Nielsen, Christer Larsson, Antonius van Maris and Jack Pronk, Current Opinion in Biotechnology 2013, 24:17 3. CellDesignerTM: A modeling tool of biochemical networks, http:\\www.celldesigner.org 4. Ana Tuek, Christian Lorenz Mller, Jochen Supper, Andreas Zell, }elimir Kurtanjek, and Ivo F. Sbalzarini: "A Systems Biology Markup Language Model of a T-Cell Receptor Activated Signal Transduction Network", Proceedings of 29-th ITI 2007 Degenring, D.:Dissertation. University of Rostock, 2004. 6. Ana Tuaek, }. Kurtanjek,  Model and global sensitivity analysis of E. coli central metabolism , Proceedings MATHMOD 09 Vienna, Austria, 2009. 7. }elimir Kurtanjek,  Probabilistic Metabolic control Analysis , Systems Biology of Microorganisms, 22-24, March, 2010, Paris, France 8. Alvarez-Vasquez, F., Riezman, H., Hannun, Y.A., Voit, E.O. Mathematical Modeling and Validation of the Ergosterol Pathway in Saccharomyces cerevisiae. PloS ONE. 6, 1-17. (2011). 9. 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