Here Is What It Looks Like

Branch-N-glycan

Designed by Ken S. Lau and James W. Dennis

Overview

The Golgi pathway trims and remodels N-glycans occupying NXS/T sites on newly synthesized glycoproteins destined for the cell surface or secretion. N-glycans are substituted with GlcNAc by multiple N-acetylglucosaminyltransferases (Mgat gene products) in the medial Golgi compartment. These enzymes initiate branches that are extended by a number of other enzymes to generate ligands for mammalian lectins. These interactions serve to localize glycoproteins to different compartments in the cell. As a major feature of this model, the Mgat enzymes form a pathway with the intriguing property of multistep ultrasensitivity to the common substrate UDP-GlcNAc. The pathway has been modeled based on 14 N-glycan structure including high mannose precursors, GlcNAc-branched intermediates with 0 to 4 branches, the bisecting GlcNAc, and extension of the branches with galactose (Fig 1). The “Branch-N-glycan” model was used to simulate cell surface N-glycan compositions for different concentrations of UDP-GlcNAc, and the results were used in a simple model for galectin binding and retention of glycoprotein receptors at the cell surface (“Lattice” model) as described in Lau et al. Cell 129:123,2007

Instructions

The content of the model as found in the “lisp” file is shown below.

It contains (1) The abbreviated names of all N-glycan structures used in the model, (2) the chemical equations specifying all the reactions and (3) rate constants and initial concentrations.

Constants can be modified as required, or the model can be used as a starting point for the addition of reactions and more structures.

To get start

Download golgomodeln.lisp

Download littleB

littleb creator is Aneil Mallavarapu and more information can be found at

Open the lisp file in the littleB program. Then press "play".

Matlab files consisting of the differential equations are generated in the same directory.

littleB converts the model from chemical equations to ODEs that can be run in Matlab. Now all you have to do is run Matlab using the regular ODE convention, for example [t,y] =golgimodel ([0 10000000]), which will allow the system to go to steady state.

Publications:

Lau KS, Partridge EA, Grigorian A, Silvescu CI, Reinhold VN, Demetriou M, Dennis JW. Complex N-glycan number and degree of branching cooperate to regulate cell proliferation and differentiation. Cell 129:123-34, 2007

Chen, H-L, Newton B, Beeton, C. Lau, KS, Lee S. Walker E, Grigorian A, van den Noort1 S, Yu Z, Siminovitch KA, HauserSL, Oksenberg JR, ChandyKG, Dennis JW, and DemetriouM. MGAT1 and CTLA-4 variants interact to regulate human autoimmunity. (2008) Cell, in review

Model of N-glycan branching (A) Dotted arrows represent transport through the trans Golgi to the cell surface. The primary input for Golgi modification Rti(glycan) is the high mannose precursors, while the primary output is the mature N-glycans on glycoproteins at steady state. Mature N-glycans are degraded at kturnover = 8.06 x 10-5 s-1, an estimate from the mean half-life of membrane components in the cell. With mass conservation, the rate of input equals the rate of output. The secondary input of the system is UDP-GlcNAc governed by the Golgi UDP-GlcNAc/UMP transporter, which operates near Vmax at physiological UDP-GlcNAc levels (100 μM). Abbreviations: branching N-acetylglycosaminyltransferases I to V(Mgat1 to 5), 1,3N-acetylglycosaminyltransferase (iGNT), -mannosidaseII (MII), 1,4Galactosyltransferase (GalT), * indicates UDP-GlcNAc-bound enzyme. N-glycans are abbreviated:

Core –Man4GlcNAc4GlcNAc

M6-9 – (Man1-4)-Man2(Man3) Man6(GlcNAc2Man3)-Core

M5Gn – Man2(Man1,3) Man6(GlcNAc2Man3)-Core

M’Gn – Man2(3) Man6(GlcNAc2Man3)-Core

M’’Gn – Man1,6(GlcNAc2Man3)-Core

GnGn – GlcNAc2Man6(GlcNAc1,2Man3)-Core

GnGnGn – Gn3(2’,4,2)-Core

GnGnGnGn – Gn4(6’,2’,4,2)-Core

G – galactosylated forms of Gn

(X)i – N-glycan with 1 glucosamine (GlcNAc…) extension

(X)I – N-glycan with 1 lactosamine (Gal4GlcNAc…) extension

(X)m – mature N-glycan on glycoprotein (not available for further processing)

(X)B – bisected sugar

(B) Simulations oftri-antennary N-glycans produced in mgat5-/- cells by the “normal” or assumed pathway structurein our model(Branched Pathway), or by a system with a single compartment no transit and no product inhibition (Linear Pathway). Simulations were run with and without mgat1 overexpression, and performed using exactly the same kinetic parameters. The accompanying table summarizes the response metrics (nH, D50, maximal fraction) of the above simulations. As noted in the model description above, product inhibition was included to simulate the intermediate and product level changes observed experimentally by mass spectrometry. Simulations performed on the removal of product inhibition or the removal of transit separately showed that either one of these factors alone is sufficient to produce ultrasensitive upregulation of tri-antennary N-glycans by UDP-GlcNAc and suppression by mgat1. From Lau et al Cell 129:123-34, 2007

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(in-package :b-user)

(include b-user/ode-biochem)

;; SPECIES

(def-species-types compartment Mgat1 UDP_GlcNAc Mgat1* M6_9+ Mgat1*M6_9+

Mgat1*M5Gn+ UDP M5Gn+ MII MIIM5Gn+ MIIM^Gn+ M^Gn+ MIIM^Gn+2 MIIM^^Gn+ M^^Gn+

Mgat2 Mgat2* Mgat2*M^^Gn+ Mgat2*GnGn+ GnGn+ Mgat4 Mgat4* Mgat4*GnGn+

Mgat4*GnGnGn+ GnGnGn+ Mgat5 Mgat5* Mgat5*GnGnGn+ Mgat5*GnGnGnGn+ GnGnGnGn+ Mgat3

Mgat3* Mgat3*GnGn+ GnGnB+ Mgat3*GnGnGn+ GnGnGnB+ Mgat3*GnGnGnGn+ GnGnGnGnB+ M5Gn

GnGn GnGnGn GnGnGnGn GnGnB GnGnGnB GnGnGnGnB GalT GalTGnGnB GGnB GalTGGnB GGB

GnGnBm GGnBm GGBm GalTM5Gn M5G M5Gnm M5Gm GalTGnGnGnB GGnGnB GalTGGnGnB GGGnB

GalTGGGnB GGGB GnGnGnBm GGnGnBm GGGnBm GGGBm GalTGnGnGnGnB GGnGnGnB GalTGGnGnGnB

GGGnGnB GalTGGGnGnB GGGGnB GalTGGGGnB GGGGB GnGnGnGnBm GGnGnGnBm GGGnGnBm

GGGGnBm GGGGBm M6_9m GalTGnGn GGn GalTGGn GG GnGnm GGnm GGm GalTGnGnGn GGnGn

GalTGGnGn GGGn GalTGGGn GGG GnGnGnm GGnGnm GGGnm GGGm GalTGnGnGnGn GGnGnGn

GalTGGnGnGn GGGnGn GalTGGGnGn GGGGn GalTGGGGn GGGG GnGnGnGnm GGnGnGnm GGGnGnm

GGGGnm GGGGm iGNT iGNT* iGNT*GGGG GGGGi iGNT*GGG GGGi iGNT*GG GGi GalTGGGGi

GGGG1 GalTGGGi GGG1 GalTGGi GG1 iGNT*GGGG1 GGGG1i iGNT*GGG1 GGG1i iGNT*GG1 GG1i

GalTGGGG1i GGGG11 GalTGGG1i GGG11 GalTGG1i GG11 GGGG1m GGG1m GG1m GGGG11m GGG11m

GG11m)

;; REACTIONS

(define mgat1bindingUDPGlcNac {Mgat1 + UDP_GlcNAc -> Mgat1*})

(define mgat1unbindingUDPGlcNac {Mgat1 + UDP_GlcNAc <- Mgat1*})

(define mgat1*bindingM6_9+ {Mgat1* + M6_9+ -> Mgat1*M6_9+})

(define mgat1*unbindingM6_9+ {Mgat1* + M6_9+ <- Mgat1*M6_9+})

(define mgat1transition {Mgat1*M6_9+ -> Mgat1*M5Gn+})

(define mgat1product {Mgat1*M5Gn+ -> UDP + Mgat1 + M5Gn+})

(define mgat1rev {Mgat1*M5Gn+ <- UDP + Mgat1 + M5Gn+})

(define maniibindingM5Gn+ {MII + M5Gn+ -> MIIM5Gn+})

(define maniiunbindingM5Gn+ {MII + M5Gn+ <- MIIM5Gn+})

(define maniitransition {MIIM5Gn+ -> MIIM^Gn+})

(define maniiproduct {MIIM^Gn+ -> MII + M^Gn+})

(define maniirev {MIIM^Gn+ <- MII + M^Gn+})

(define maniibindingM^Gn+ {MII + M^Gn+ -> MIIM^Gn+2})

(define maniiunbindingM^Gn+ {MII + M^Gn+ <- MIIM^Gn+2})

(define maniitransition2 {MIIM^Gn+2 -> MIIM^^Gn+})

(define maniiproduct2 {MIIM^^Gn+ -> MII + M^^Gn+})

(define maniirev2 {MIIM^^Gn+ <- MII + M^^Gn+})

(define mgat2bindingUDPGlcNac {Mgat2 + UDP_GlcNAc -> Mgat2*})

(define mgat2unbindingUDPGlcNac {Mgat2 + UDP_GlcNAc <- Mgat2*})

(define mgat2*bindingM^^Gn+ {Mgat2* + M^^Gn+ -> Mgat2*M^^Gn+})

(define mgat2*unbindingM^^Gn+ {Mgat2* + M^^Gn+ <- Mgat2*M^^Gn+})

(define mgat2transition {Mgat2*M^^Gn+ -> Mgat2*GnGn+})

(define mgat2product {Mgat2*GnGn+ -> UDP + Mgat2 + GnGn+})

(define mgat2rev {Mgat2*GnGn+ <- UDP + Mgat2 + GnGn+})

(define mgat4bindingUDPGlcNac {Mgat4 + UDP_GlcNAc -> Mgat4*})

(define mgat4unbindingUDPGlcNac {Mgat4 + UDP_GlcNAc <- Mgat4*})

(define mgat4*bindingGnGn+ {Mgat4* + GnGn+ -> Mgat4*GnGn+})

(define mgat4*unbindingGnGn+ {Mgat4* + GnGn+ <- Mgat4*GnGn+})

(define mgat4transition {Mgat4*GnGn+ -> Mgat4*GnGnGn+})

(define mgat4product {Mgat4*GnGnGn+ -> UDP + Mgat4 + GnGnGn+})

(define mgat4rev {Mgat4*GnGnGn+ <- UDP + Mgat4 + GnGnGn+})

(define mgat5bindingUDPGlcNac {Mgat5 + UDP_GlcNAc -> Mgat5*})

(define mgat5unbindingUDPGlcNac {Mgat5 + UDP_GlcNAc <- Mgat5*})

(define mgat5*bindingGnGnGn+ {Mgat5* + GnGnGn+ -> Mgat5*GnGnGn+})

(define mgat5*unbindingGnGnGn+ {Mgat5* + GnGnGn+ <- Mgat5*GnGnGn+})

(define mgat5transition {Mgat5*GnGnGn+ -> Mgat5*GnGnGnGn+})

(define mgat5product {Mgat5*GnGnGnGn+ -> UDP + Mgat5 + GnGnGnGn+})

(define mgat5rev {Mgat5*GnGnGnGn+ <- UDP + Mgat5 + GnGnGnGn+})

(define mgat3bindingUDPGlcNac {Mgat3 + UDP_GlcNAc -> Mgat3*})

(define mgat3unbindingUDPGlcNac {Mgat3 + UDP_GlcNAc <- Mgat3*})

(define mgat3*bindingGnGn+ {Mgat3* + GnGn+ -> Mgat3*GnGn+})

(define mgat3*unbindingGnGn+ {Mgat3* + GnGn+ <- Mgat3*GnGn+})

(define mgat3product2 {Mgat3*GnGn+ -> UDP + Mgat3 + GnGnB+})

(define mgat3*bindingGnGnGn+ {Mgat3* + GnGnGn+ -> Mgat3*GnGnGn+})

(define mgat3*unbindingGnGnGn+ {Mgat3* + GnGnGn+ <- Mgat3*GnGnGn+})

(define mgat3product3 {Mgat3*GnGnGn+ -> UDP + Mgat3 + GnGnGnB+})

(define mgat3*bindingGnGnGnGn+ {Mgat3* + GnGnGnGn+ -> Mgat3*GnGnGnGn+})

(define mgat3*unbindingGnGnGnGn+ {Mgat3* + GnGnGnGn+ <- Mgat3*GnGnGnGn+})

(define mgat3product4 {Mgat3*GnGnGnGn+ -> UDP + Mgat3 + GnGnGnGnB+})

(define M5Gntransit {M5Gn+ -> M5Gn})

(define GnGntransit {GnGn+ -> GnGn})

(define GnGnGntransit {GnGnGn+ -> GnGnGn})

(define GnGnGnGntransit {GnGnGnGn+ -> GnGnGnGn})

(define GnGnBtransit {GnGnB+ -> GnGnB})

(define GnGnGnBtransit {GnGnGnB+ -> GnGnGnB})

(define GnGnGnGnBtransit {GnGnGnGnB+ -> GnGnGnGnB})

(define GalTbindingGnGnB {GnGnB + GalT -> GalTGnGnB})

(define GalTunbindingGnGnB {GnGnB + GalT <- GalTGnGnB})

(define GalTGnGnBproduct {GalTGnGnB -> GGnB + GalT})

(define GalTbindingGGnB {GGnB + GalT -> GalTGGnB})

(define GalTunbindingGGnB {GGnB + GalT <- GalTGGnB})

(define GalTGGnBproduct {GalTGGnB -> GGB + GalT})

(define GnGnBmature {GnGnB -> GnGnBm})

(define GGnBmature {GGnB -> GGnBm})

(define GGBmature {GGB -> GGBm})

(define GalTbindingM5Gn {M5Gn + GalT -> GalTM5Gn})

(define GalTunbindingM5Gn {M5Gn + GalT <- GalTM5Gn})

(define GalTM5Gnproduct {GalTM5Gn -> M5G + GalT})

(define M5Gnmature {M5Gn -> M5Gnm})

(define M5Gmature {M5G -> M5Gm})

(define GalTbindingGnGnGnB {GnGnGnB + GalT -> GalTGnGnGnB})

(define GalTunbindingGnGnGnB {GnGnGnB + GalT <- GalTGnGnGnB})

(define GalTGnGnGnBproduct {GalTGnGnGnB -> GGnGnB + GalT})

(define GalTbindingGGnGnB {GGnGnB + GalT -> GalTGGnGnB})

(define GalTunbindingGGnGnB {GGnGnB + GalT <- GalTGGnGnB})

(define GalTGGnGnBproduct {GalTGGnGnB -> GGGnB + GalT})

(define GalTbindingGGGnB {GGGnB + GalT -> GalTGGGnB})

(define GalTunbindingGGGnB {GGGnB + GalT <- GalTGGGnB})

(define GalTGGGnBproduct {GalTGGGnB -> GGGB + GalT})

(define GnGnGnBmature {GnGnGnB -> GnGnGnBm})

(define GGnGnBmature {GGnGnB -> GGnGnBm})

(define GGGnBmature {GGGnB -> GGGnBm})

(define GGGBmature {GGGB -> GGGBm})

(define GalTbindingGnGnGnGnB {GnGnGnGnB + GalT -> GalTGnGnGnGnB})

(define GalTunbindingGnGnGnGnB {GnGnGnGnB + GalT <- GalTGnGnGnGnB})

(define GalTGnGnGnGnBproduct {GalTGnGnGnGnB -> GGnGnGnB + GalT})

(define GalTbindingGGnGnGnB {GGnGnGnB + GalT -> GalTGGnGnGnB})

(define GalTunbindingGGnGnGnB {GGnGnGnB + GalT <- GalTGGnGnGnB})

(define GalTGGnGnGnBproduct {GalTGGnGnGnB -> GGGnGnB + GalT})

(define GalTbindingGGGnGnB {GGGnGnB + GalT -> GalTGGGnGnB})

(define GalTunbindingGGGnGnB {GGGnGnB + GalT <- GalTGGGnGnB})

(define GalTGGGnGnBproduct {GalTGGGnGnB -> GGGGnB + GalT})

(define GalTbindingGGGGnB {GGGGnB + GalT -> GalTGGGGnB})

(define GalTunbindingGGGGnB {GGGGnB + GalT <- GalTGGGGnB})

(define GalTGGGGnBproduct {GalTGGGGnB -> GGGGB + GalT})

(define GnGnGnGnBmature {GnGnGnGnB -> GnGnGnGnBm})

(define GGnGnGnBmature {GGnGnGnB -> GGnGnGnBm})

(define GGGnGnBmature {GGGnGnB -> GGGnGnBm})

(define GGGGnBmature {GGGGnB -> GGGGnBm})

(define GGGGBmature {GGGGB -> GGGGBm})

(define M6_9mature {M6_9+ -> M6_9m})

(define GalTbindingGnGn {GnGn + GalT -> GalTGnGn})

(define GalTunbindingGnGn {GnGn + GalT <- GalTGnGn})

(define GalTGnGnproduct {GalTGnGn -> GGn + GalT})

(define GalTbindingGGn {GGn + GalT -> GalTGGn})

(define GalTunbindingGGn {GGn + GalT <- GalTGGn})

(define GalTGGnproduct {GalTGGn -> GG + GalT})

(define GnGnmature {GnGn -> GnGnm})

(define GGnmature {GGn -> GGnm})

(define GGmature {GG -> GGm})

(define GalTbindingGnGnGn {GnGnGn + GalT -> GalTGnGnGn})

(define GalTunbindingGnGnGn {GnGnGn + GalT <- GalTGnGnGn})

(define GalTGnGnGnproduct {GalTGnGnGn -> GGnGn + GalT})

(define GalTbindingGGnGn {GGnGn + GalT -> GalTGGnGn})

(define GalTunbindingGGnGn {GGnGn + GalT <- GalTGGnGn})

(define GalTGGnGnproduct {GalTGGnGn -> GGGn + GalT})

(define GalTbindingGGGn {GGGn + GalT -> GalTGGGn})

(define GalTunbindingGGGn {GGGn + GalT <- GalTGGGn})

(define GalTGGGnproduct {GalTGGGn -> GGG + GalT})

(define GnGnGnmature {GnGnGn -> GnGnGnm})

(define GGnGnmature {GGnGn -> GGnGnm})

(define GGGnmature {GGGn -> GGGnm})

(define GGGmature {GGG -> GGGm})

(define GalTbindingGnGnGnGn {GnGnGnGn + GalT -> GalTGnGnGnGn})

(define GalTunbindingGnGnGnGn {GnGnGnGn + GalT <- GalTGnGnGnGn})

(define GalTGnGnGnGnproduct {GalTGnGnGnGn -> GGnGnGn + GalT})

(define GalTbindingGGnGnGn {GGnGnGn + GalT -> GalTGGnGnGn})

(define GalTunbindingGGnGnGn {GGnGnGn + GalT <- GalTGGnGnGn})

(define GalTGGnGnGnproduct {GalTGGnGnGn -> GGGnGn + GalT})

(define GalTbindingGGGnGn {GGGnGn + GalT -> GalTGGGnGn})

(define GalTunbindingGGGnGn {GGGnGn + GalT <- GalTGGGnGn})

(define GalTGGGnGnproduct {GalTGGGnGn -> GGGGn + GalT})

(define GalTbindingGGGGn {GGGGn + GalT -> GalTGGGGn})

(define GalTunbindingGGGGn {GGGGn + GalT <- GalTGGGGn})

(define GalTGGGGnproduct {GalTGGGGn -> GGGG + GalT})

(define GnGnGnGnmature {GnGnGnGn -> GnGnGnGnm})

(define GGnGnGnmature {GGnGnGn -> GGnGnGnm})

(define GGGnGnmature {GGGnGn -> GGGnGnm})

(define GGGGnmature {GGGGn -> GGGGnm})

(define GGGGmature {GGGG -> GGGGm})

(define iGNTbindingUDPGlcNac {iGNT + UDP_GlcNAc -> iGNT*})

(define iGNTunbindingUDPGlcNac {iGNT + UDP_GlcNAc <- iGNT*})

(define iGNT*bindingGGGG {iGNT* + GGGG -> iGNT*GGGG})

(define iGNT*unbindingGGGG {iGNT* + GGGG <- iGNT*GGGG})

(define iGNT*GGGGproduct {iGNT*GGGG -> GGGGi + iGNT + UDP})

(define iGNT*bindingGGG {iGNT* + GGG -> iGNT*GGG})

(define iGNT*unbindingGGG {iGNT* + GGG <- iGNT*GGG})

(define iGNT*GGGproduct {iGNT*GGG -> GGGi + iGNT + UDP})

(define iGNT*bindingGG {iGNT* + GG -> iGNT*GG})

(define iGNT*unbindingGG {iGNT* + GG <- iGNT*GG})

(define iGNT*GGproduct {iGNT*GG -> GGi + iGNT + UDP})

(define GalTbindingGGGGi {GGGGi + GalT -> GalTGGGGi})

(define GalTunbindingGGGGi {GGGGi + GalT <- GalTGGGGi})

(define GalTGGGGiproduct {GalTGGGGi -> GGGG1 + GalT})

(define GalTbindingGGGi {GGGi + GalT -> GalTGGGi})

(define GalTunbindingGGGi {GGGi + GalT <- GalTGGGi})

(define GalTGGGiproduct {GalTGGGi -> GGG1 + GalT})

(define GalTbindingGGi {GGi + GalT -> GalTGGi})

(define GalTunbindingGGi {GGi + GalT <- GalTGGi})

(define GalTGGiproduct {GalTGGi -> GG1 + GalT})

(define iGNT*bindingGGGG1 {iGNT* + GGGG1 -> iGNT*GGGG1})

(define iGNT*unbindingGGGG1 {iGNT* + GGGG1 <- iGNT*GGGG1})

(define iGNT*GGGG1product {iGNT*GGGG1 -> GGGG1i + iGNT + UDP})

(define iGNT*bindingGGG1 {iGNT* + GGG1 -> iGNT*GGG1})

(define iGNT*unbindingGGG1 {iGNT* + GGG1 <- iGNT*GGG1})

(define iGNT*GGG1product {iGNT*GGG1 -> GGG1i + iGNT + UDP})

(define iGNT*bindingGG1 {iGNT* + GG1 -> iGNT*GG1})

(define iGNT*unbindingGG1 {iGNT* + GG1 <- iGNT*GG1})

(define iGNT*GG1product {iGNT*GG1 -> GG1i + iGNT + UDP})

(define GalTbindingGGGG1i {GGGG1i + GalT -> GalTGGGG1i})

(define GalTunbindingGGGG1i {GGGG1i + GalT <- GalTGGGG1i})

(define GalTGGGG1iproduct {GalTGGGG1i -> GGGG11 + GalT})

(define GalTbindingGGG1i {GGG1i + GalT -> GalTGGG1i})

(define GalTunbindingGGG1i {GGG1i + GalT <- GalTGGG1i})

(define GalTGGG1iproduct {GalTGGG1i -> GGG11 + GalT})

(define GalTbindingGG1i {GG1i + GalT -> GalTGG1i})

(define GalTunbindingGG1i {GG1i + GalT <- GalTGG1i})

(define GalTGG1iproduct {GalTGG1i -> GG11 + GalT})

(define GGGG1mature {GGGG1 -> GGGG1m})

(define GGG1mature {GGG1 -> GGG1m})

(define GG1mature {GG1 -> GG1m})

(define GGGG11mature {GGGG11 -> GGGG11m})

(define GGG11mature {GGG11 -> GGG11m})

(define GG11mature {GG11 -> GG11m})

(define UDPGlcNAcfwd {UDP -> UDP_GlcNAc})

(define M6_9mdegrade {M6_9m -> M6_9+})

(define M5Gnmdegrade {M5Gnm -> M6_9+})

(define M5Gmdegrade {M5Gm -> M6_9+})

(define GnGnmdegrade {GnGnm -> M6_9+})

(define GGnmdegrade {GGnm -> M6_9+})

(define GGmdegrade {GGm -> M6_9+})

(define GnGnGnmdegrade {GnGnGnm -> M6_9+})

(define GGnGnmdegrade {GGnGnm -> M6_9+})

(define GGGnmdegrade {GGGnm -> M6_9+})

(define GGGmdegrade {GGGm -> M6_9+})

(define GnGnGnGnmdegrade {GnGnGnGnm -> M6_9+})

(define GGnGnGnmdegrade {GGnGnGnm -> M6_9+})

(define GGGnGnmdegrade {GGGnGnm -> M6_9+})

(define GGGGnmdegrade {GGGGnm -> M6_9+})

(define GGGGmdegrade {GGGGm -> M6_9+})

(define GnGnBmdegrade {GnGnBm -> M6_9+})

(define GGnBmdegrade {GGnBm -> M6_9+})

(define GGBmdegrade {GGBm -> M6_9+})

(define GnGnGnBmdegrade {GnGnGnBm -> M6_9+})

(define GGnGnBmdegrade {GGnGnBm -> M6_9+})

(define GGGnBmdegrade {GGGnBm -> M6_9+})

(define GGGBmdegrade {GGGBm -> M6_9+})

(define GnGnGnGnBmdegrade {GnGnGnGnBm -> M6_9+})

(define GGnGnGnBmdegrade {GGnGnGnBm -> M6_9+})

(define GGGnGnBmdegrade {GGGnGnBm -> M6_9+})

(define GGGGnBmdegrade {GGGGnBm -> M6_9+})

(define GGGGBmdegrade {GGGGBm -> M6_9+})

(define GG1mdegrade {GG1m -> M6_9+})

(define GG11mdegrade {GG11m -> M6_9+})

(define GGG1mdegrade {GGG1m -> M6_9+})

(define GGG11mdegrade {GGG11m -> M6_9+})

(define GGGG1mdegrade {GGGG1m -> M6_9+})

(define GGGG11mdegrade {GGGG11m -> M6_9+})

;; RATE CONSTANTS

mgat1bindingUDPGlcNac.(set-rate-function 'mass-action 1.7994e+006)

mgat1unbindingUDPGlcNac.(set-rate-function 'mass-action 55.2788)

mgat1*bindingM6_9+.(set-rate-function 'mass-action 34039)

mgat1*unbindingM6_9+.(set-rate-function 'mass-action 55.2788)

mgat1transition.(set-rate-function 'mass-action 13.8197)

mgat1product.(set-rate-function 'mass-action 10)

mgat1rev.(set-rate-function 'mass-action 1.5)

maniibindingM5Gn+.(set-rate-function 'mass-action 518.8667)

maniiunbindingM5Gn+.(set-rate-function 'mass-action 12.4528)

maniitransition.(set-rate-function 'mass-action 3.1132)

maniiproduct.(set-rate-function 'mass-action 10)

maniirev.(set-rate-function 'mass-action 1.5)

maniibindingM^Gn+.(set-rate-function 'mass-action 1.0377e+003)

maniiunbindingM^Gn+.(set-rate-function 'mass-action 12.4528)

maniitransition2.(set-rate-function 'mass-action 3.1132)

maniiproduct2.(set-rate-function 'mass-action 10)

maniirev2.(set-rate-function 'mass-action 1.5)

mgat2bindingUDPGlcNac.(set-rate-function 'mass-action 123080)

mgat2unbindingUDPGlcNac.(set-rate-function 'mass-action 94.5268)

mgat2*bindingM^^Gn+.(set-rate-function 'mass-action 621890)

mgat2*unbindingM^^Gn+.(set-rate-function 'mass-action 94.5268)

mgat2transition.(set-rate-function 'mass-action 23.6317)

mgat2product.(set-rate-function 'mass-action 10)

mgat2rev.(set-rate-function 'mass-action 1.5)

mgat4bindingUDPGlcNac.(set-rate-function 'mass-action 1.6892e+003)

mgat4unbindingUDPGlcNac.(set-rate-function 'mass-action 6.7588)

mgat4*bindingGnGn+.(set-rate-function 'mass-action 1.1570e+004)

mgat4*unbindingGnGn+.(set-rate-function 'mass-action 6.7588)

mgat4transition.(set-rate-function 'mass-action 1.6892)

mgat4product.(set-rate-function 'mass-action 10)

mgat4rev.(set-rate-function 'mass-action 0.15)

mgat5bindingUDPGlcNac.(set-rate-function 'mass-action 12045.45)

mgat5unbindingUDPGlcNac.(set-rate-function 'mass-action 106)

mgat5*bindingGnGnGn+.(set-rate-function 'mass-action 1522988.5)

mgat5*unbindingGnGnGn+.(set-rate-function 'mass-action 106)

mgat5transition.(set-rate-function 'mass-action 26.5)

mgat5product.(set-rate-function 'mass-action 10)

mgat5rev.(set-rate-function 'mass-action 0.075)

mgat3bindingUDPGlcNac.(set-rate-function 'mass-action 6057.14)

mgat3unbindingUDPGlcNac.(set-rate-function 'mass-action 2.0352)

mgat3*bindingGnGn+.(set-rate-function 'mass-action 121142.86)

mgat3*unbindingGnGn+.(set-rate-function 'mass-action 2.0352)

mgat3product2.(set-rate-function 'mass-action 0.5088)

mgat3*bindingGnGnGn+.(set-rate-function 'mass-action 121142.86)

mgat3*unbindingGnGnGn+.(set-rate-function 'mass-action 2.0352)

mgat3product3.(set-rate-function 'mass-action 0.5088)

mgat3*bindingGnGnGnGn+.(set-rate-function 'mass-action 121142.86)

mgat3*unbindingGnGnGnGn+.(set-rate-function 'mass-action 2.0352)

mgat3product4.(set-rate-function 'mass-action 0.5088)

M5Gntransit.(set-rate-function 'mass-action 0.003)

GnGntransit.(set-rate-function 'mass-action 0.003)

GnGnGntransit.(set-rate-function 'mass-action 0.003)

GnGnGnGntransit.(set-rate-function 'mass-action 0.003)

GnGnBtransit.(set-rate-function 'mass-action 0.003)

GnGnGnBtransit.(set-rate-function 'mass-action 0.003)

GnGnGnGnBtransit.(set-rate-function 'mass-action 0.003)

GalTbindingGnGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGnGnB.(set-rate-function 'mass-action 57.9876)

GalTGnGnBproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGnB.(set-rate-function 'mass-action 57.9876)

GalTGGnBproduct.(set-rate-function 'mass-action 14.4969)

GnGnBmature.(set-rate-function 'mass-action 0.003)

GGnBmature.(set-rate-function 'mass-action 0.003)

GGBmature.(set-rate-function 'mass-action 0.003)

GalTbindingM5Gn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingM5Gn.(set-rate-function 'mass-action 57.9876)

GalTM5Gnproduct.(set-rate-function 'mass-action 14.4969)

M5Gnmature.(set-rate-function 'mass-action 0.003)

M5Gmature.(set-rate-function 'mass-action 0.003)

GalTbindingGnGnGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGnGnGnB.(set-rate-function 'mass-action 57.9876)

GalTGnGnGnBproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGnGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGnGnB.(set-rate-function 'mass-action 57.9876)

GalTGGnGnBproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGGnB.(set-rate-function 'mass-action 57.9876)

GalTGGGnBproduct.(set-rate-function 'mass-action 14.4969)

GnGnGnBmature.(set-rate-function 'mass-action 0.003)

GGnGnBmature.(set-rate-function 'mass-action 0.003)

GGGnBmature.(set-rate-function 'mass-action 0.003)

GGGBmature.(set-rate-function 'mass-action 0.003)

GalTbindingGnGnGnGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGnGnGnGnB.(set-rate-function 'mass-action 57.9876)

GalTGnGnGnGnBproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGnGnGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGnGnGnB.(set-rate-function 'mass-action 57.9876)

GalTGGnGnGnBproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGGnGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGGnGnB.(set-rate-function 'mass-action 57.9876)

GalTGGGnGnBproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGGGnB.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGGGnB.(set-rate-function 'mass-action 57.9876)

GalTGGGGnBproduct.(set-rate-function 'mass-action 14.4969)

GnGnGnGnBmature.(set-rate-function 'mass-action 0.003)

GGnGnGnBmature.(set-rate-function 'mass-action 0.003)

GGGnGnBmature.(set-rate-function 'mass-action 0.003)

GGGGnBmature.(set-rate-function 'mass-action 0.003)

GGGGBmature.(set-rate-function 'mass-action 0.003)

M6_9mature.(set-rate-function 'mass-action 5)

GalTbindingGnGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGnGn.(set-rate-function 'mass-action 57.9876)

GalTGnGnproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGn.(set-rate-function 'mass-action 57.9876)

GalTGGnproduct.(set-rate-function 'mass-action 14.4969)

GnGnmature.(set-rate-function 'mass-action 0.003)

GGnmature.(set-rate-function 'mass-action 0.003)

GGmature.(set-rate-function 'mass-action 0.003)

GalTbindingGnGnGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGnGnGn.(set-rate-function 'mass-action 57.9876)

GalTGnGnGnproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGnGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGnGn.(set-rate-function 'mass-action 57.9876)

GalTGGnGnproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGGn.(set-rate-function 'mass-action 57.9876)

GalTGGGnproduct.(set-rate-function 'mass-action 14.4969)

GnGnGnmature.(set-rate-function 'mass-action 0.003)

GGnGnmature.(set-rate-function 'mass-action 0.003)

GGGnmature.(set-rate-function 'mass-action 0.003)

GGGmature.(set-rate-function 'mass-action 0.003)

GalTbindingGnGnGnGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGnGnGnGn.(set-rate-function 'mass-action 57.9876)

GalTGnGnGnGnproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGnGnGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGnGnGn.(set-rate-function 'mass-action 57.9876)

GalTGGnGnGnproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGGnGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGGnGn.(set-rate-function 'mass-action 57.9876)

GalTGGGnGnproduct.(set-rate-function 'mass-action 14.4969)

GalTbindingGGGGn.(set-rate-function 'mass-action 3.8150e+004)

GalTunbindingGGGGn.(set-rate-function 'mass-action 57.9876)

GalTGGGGnproduct.(set-rate-function 'mass-action 14.4969)

GnGnGnGnmature.(set-rate-function 'mass-action 0.003)