ABSTRACT
Phosphate limited grown Anabaena variabilis has the capability of processing information about external phosphate fluctuations by means of interconnected adaptive events. Adaptive events are physiological processes that are characterized by two opposite manifestations, namely adapted states and adaptive operation modes. In adapted states the energy-converting constituents of the uptake system operate under the prevailing external conditions in a coherent manner with least energy dissipation. Adaptive operation modes take place when adapted states are disturbed by persistent changes in phosphate supply. In this mode the outcome of former adaptations to elevated phosphate levels guides the emergence of a new adapted state. The influence of antecedent adapted states on subsequent adaptations was studied experimentally and characteristic examples for such information processing are given. The theory of self-referential systems allowed analyzing these examples. For this purpose adaptive events had to be considered as elements of a communicating network, in which, along a historic succession of alternating adapted states and adaptive operation modes, information pertaining to the self-preservation of the organism is transferred from one adaptive event to the next: the latter "interprets" environmental changes by means of distinct adaptive operation modes, aimed at preservation of the organism. The result of this interpretation is again leading to a coherent state that is passed on to subsequent adaptive events. A generalization of this idea to the adaptive interplay of other energy converting subsystems of the cell leads to the dynamic view of cellular information processing in which the organism recreates itself in every new experience.
ABSTRACT
We present a model of microbial information processing that contains characteristic features of the phenomenon of physiological adaptation. The backbone of the model is the "adaptive event" in which energy-converting subsystems of the cell interact with the changing environment. In this process, the subsystems pass, via an adaptive operation mode, from one adapted state to the next. An adaptive operation mode takes place when an adapted state is disturbed by an environmental alteration. These two manifestations of an adaptive event were differently treated in the simulation, based on an application of linear irreversible thermodynamics to the energy transduction of adaptive subsystems. In adapted states, the conductivity coefficients of the flow-force relationships employed remained constant, whereas during an adaptive operation mode, these coefficients were altered in a directional manner during the simulation. An example dealing with the complex relationship between phosphate uptake and cyanobacterial growth is given. In this example, the simulation of adapted states of two subsystems of the incorporating machinery, namely the phosphate carrier in the cell membrane and the F-ATPase in the thylakoid membrane, was in accordance with the measured uptake kinetics, and when fixed, predetermined conductivity coefficients were used. In the adaptive operation mode, however, the simulated behavior was in agreement with experimental observations when the program was able to "interpret" its own performance in the light of environmental phosphate fluctuations, experienced by the cell in the past, and to reconstruct the two subsystems according to this interpretation. Via transitions between adapted states and adaptive modes, information is transferred from one adaptive event to the next: the latter "inherits" the results of former interpretations. By appropriating them selectively, it is entering into a future in which its own interpretation is passed on to the following adaptive event. The model is discussed with respect to the concept of autopoiesis.
