Biased pseudpods Sample Clauses

Biased pseudpods. ‌ The models listed so far are compass based models. They are based on the proposition that signaling precedes the generation of well placed pseudopods. These models are a natural extension of the prevailing "gradient sensing => polarisation => movement" dogma. Several recent observations however conflict with this proposition: i; chemo- taxis at low gradients is best described by a biased random walk. ii; Pseudopods are extended at a constant rate irrespective of the cells orientation in a gradient. iii; Un- favourable pseudopods can be retracted. iv; New pseudopods originate mostly from previous ones [6, 1]. Observations i and ii show that D. Discoideum cells move by default and generate pseudpods at a constant speed. In order to move directionally, regulation should take place not on when the pseudopods are generated but on where. This observation agrees with the finding that the Ras/PI3K/F-actin system does not require G protein input [80] but instead facilitates polarization leading to random movement in the absence of cAMP. When a gradient is applied, this autonomous sys- tem receives directional input and polarizes in the correct direction. Observation iii and iv suggest that not only do cells show persistence in their trajectories because of the fact that new pseudopods are (mostly) restricted to the current leading edge, a form of temporal sampling also plays a role and the decision to keep a pseudopod is made after its generation. This mechanism is reminiscent of bacterial chemotaxis which functions by a higher persistence in "correct" directions [92]. Taken together this leads to a model where instead of the gradient determining the correct direction for a pseudopod, pseudopods are positioned with a certain probability around the cell. The input parameters that govern pseudopod positioning are gradient steepness, direction and the position of the previous pseudopod. Such a model implies; i; a steeper gradient will lead to a higher directional accuracy due to more pseudopods being placed in the correct direction, ii; deviation of the cells polarity axis with re- spect to the gradient will lead to a corresponding bias in the probability distribution of pseudopod generation and iii; dependence of pseudopod position on the position of the previous pseudopod will lead to autocorrelation in the cells movement charac- terized by a certain persistence time. Indeed, the observation of thousands of cells reveals the probabilistic nature of pseudopod gener...
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