A physiological analysis of walking in the American lobster (Homarus americanus).

The normal, unrestrained, forward walking of the lobster was studied with a closed-circuit television system and a video-tape recorder. A frame-by-frame analysis was undertaken and measurements made of unilateral stepping sequences, contralateral and ipsilateral phase relations between pairs of legs, the movements at the leg joints primarily involved in stepping and their differences in each of the four pereiopods. The order of stepping was expressed in terms of the probability of any leg following any other leg and it was found that while there is a preferred order, there is considerable variation from the dominant pattern. The commonest deviations from the dominant gait are those involving the simplest types of re-ordering of the sequence. Pairs of contralateral legs show a strong tendency to alternate but all phase relations can occur. Similarly, while the ipsilateral legs show preferred phase relations, all possible relations do occur. The four pereiopods from anterior to posterior were found to have respectively, a pulling action, a combined pulling and rowing action, a rowing action and a combined pushing and rowing action. The same parameters of stepping were recorded from animals walking on a transparent, driven treadmill and, as no significant differences were found in comparisons with results from freely moving animals, subsequent results were obtained from animals walking on the treadmill where more detailed study and manipulation could readily be made. The stepping action of the third pereiopod during forwards walking involves major movements about two joints whereas the other pereiopods move about three joints. Detailed study of the intra-leg activity was therefore confined to the third pereiopod where the simpler action considerably simplified the problems involved in collecting and analysing data. Measurements were made of the angles swept out by the joints of the third pereiopod during movement. Electromyograms were recorded from the six muscles primarily responsible for these movements and from movement transducers placed at the joints. The duration of the movements and of the bursts of acitivity in the muscles and the interrelation between different muscle bursts were measured and a computer-aided analysis made to determine the characteristic features of the inter-burst relations during stepping. While there is considerable variability from step to step, the overall activity is relatively phase-constant over a wide range of stepping frequencies. When some of the key parameters of normal walking had been characterized, changes designed to alter the sensory input to the system were imposed.