V Shalve h substrate V Shalve h substrate h substrate Shalve V h V substrate h Shalve h substrate h compartment_1 k1 species_1 compartment_1 k1 species_2 compartment_1 k1 species_3 compartment_1 k1 species_4 compartment_1 parameter_1 species_3 compartment_1 parameter_2 species_4 compartment_1 function_1 V Shalve parameter_4 species_2 compartment_1 function_2 species_1 Shalve V parameter_5 compartment_1 parameter_6 species_4