Tarsal intersegmental reflex responses in the locust hind leg

Abstract

Locomotion is vital for vertebrates and invertebrates to survive. However, the mechanisms for locomotion are partially unknown. Central Pattern Generators and reflex systems have been shown to be the basis of most movements performed by arthropods. Much has been investigated lately on Central Pattern Generators, but little work has been done in reflex systems. Locomotion and motor output in feet (or tarsus in arthropods) has also been disregarded in research. Despite that feet are responsible for stability and agility in most animals, research on feet movements is scarce.

In this thesis the tarsal intersegmental reflex of the locust hind leg is investigated. The tarsal reflex consists of a response in the tarsus when there is a change in the femoro-tibial joint. The main objective of the thesis is to describe the system and to develop mathematical and experimental methods to study, model and analyse it. Through a set of experiments is shown that as the knee joint is extended, the tarsus is depressed, and as the knee joint flexes, the tarsus levates. The experiments demonstrated that there is a purely neuronal link between the femoro-tibial joint position and the tibio-tarsal joint position. Moreover, it also reveals the effect of neuromodulatory compounds, such as dopamine, serotonin or octopamine. The tarsal reflex responses are fairly consistent across individuals, although significant variability across animals was found.

To model a system where variability is an issue, a mathematical model with strong generalisation abilities is used: Artificial Neural Networks (ANNs). To design the ANNs, a metaheuristic algorithm has been implemented. The resulting ANNs are shown to be as accurate as other mathematical models used in physiology when used in a well known reflex system, the FETi responses. This results showed that ANNs are as good as Wiener methods in predicting responses and they outperform them in prediction of Gaussian inputs. Furthermore, they are able to predict responses in different animals, independently of the variability, with a more limited performance.

New experimental methods are also designed to obtain accurate recordings of tarsal movements in response to knee joint changes. These experimental methods facilitate the data acquisition and its accuracy, reducing measurement errors. Using the mathematical methods validated, these responses are modelled and studied, showing responses to Gaussian and sinusoidal inputs, variability across individuals and effects of neuromodulators.

With the tarsal reflex described and modelled, it can be used as a tool for further research in disciplines such as medicine, in the diagnose and treatment of euromuscular dysfunction or design of prosthesis and orthoses. This model can also be implemented in robotics to aid in stability when walking on irregular terrain.

More information

Identifiers

ORCID for David Simpson: orcid.org/0000-0001-9072-5088

Catalogue record

Export record