Models for photosynthesis, part 3

Here I continue the discussion of models for photosynthesis in two previous posts. There I described the Pettersson and Poolman models and indicated the possibility of introducing variants of these which use exclusively mass action kinetics. I call these the Pettersson-MA and Poolman-MA models. I was interested in obtaining information about the qualitative behaviour of solutions of these ODE systems. This gave rise to the MSc project of Dorothea Möhring which she recently completed successfully. Now we have extended this work a little further and have written up the results in a paper which has just been uploaded to ArXiv. The central issue is that of overload breakdown which is related to the mathematical notion of persistence. We would like to know under what circumstances a positive solution can have -limit points where some concentrations vanish and, if so, which concentrations vanish in that case. It seems that there was almost no information on the latter point in the literature so that the question of what exactly overload breakdown is remained a bit nebulous. The general idea is that the Pettersson model should have a stronger tendency to undergo overload breakdown while the Poolman model should have a stronger tendency to avoid it. The Pettersson-MA and Poolman-MA models represent a simpler context to work in to start with.

For the Pettersson-MA model we were able to identify a regime in which overload breakdown takes place. This is where the initial concentrations of all sugar phosphates and inorganic phosphate in the chloroplast are sufficiently small. In that case the concentrations of all sugar phosphates tend to zero at late times with two exceptions. The concentrations of xylulose-4-phosphate and sedoheptulose-7-phosphate do not tend to zero. These results are obtained by linearizing the system around a simple stationary solution on the boundary and applying the centre manifold theorem. Another result is that if the reaction constants satisfy a certain inequality a positive solution can have no positive -limit points. In particular, there are no positive stationary solutions in that case. This is proved using a Lyapunov function related to the total number of carbon atoms. In the case of the Poolman-MA model it was shown that the stationary point which was stable in the Pettersson case becomes unstable. Moreover, a quantitative lower bound for concentration of sugar phosphates at late times in obtained.These results fit well with the intuitive picture of what should happen. Some of the results on the Poolman-MA model can be extended to analogous ones for the original Poolman model. On the other hand the task of giving a full rigorous definition of the Pettersson model was postponed for later work. The direction in which this could go has been sketched in a previous post.

There remains a lot to be done. It is possible to define a kind of hybrid model by setting in the Poolman model. It would be desirable to completely clarify the definition of the Pettersson model and then, perhaps, to show that it can be obtained as a well-behaved limiting system of the hybrid system in the sense of geometric singular perturbation theory. This might allow the dynamical properties of solutions of the different systems to be related to each other. The only result on stationary solutions obtained so far is a non-existence theorem. It would be of great interest to have positive results on the existence, multiplicity and stability of stationary solutions. A related question is that of classifying possible -limit points of positive solutions where some of the concentrations are zero. This was done in part in the paper but what was not settled is whether potential -limit points with positive concentrations of the hexose phosphates can actually occur. Finally, there are a lot of other models for the Calvin cycle on the market and it would be interesting to see to what extent they are susceptible to methods similar to those used in our paper.

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