Spinal reflexes, mechanisms and concepts: from Eccles to Lundberg and beyond.

This review focuses on investigations by Sir John Eccles and co-workers in Canberra, AUS in the 1950s, in which they used intracellular recordings to unravel the organization of neuronal networks in the cat spinal cord. Five classical spinal reflexes are emphasized: recurrent inhibition of motoneurons via motor axon collaterals and Renshaw cells, pathways from muscle spindles and Golgi tendon organs, presynaptic inhibition, and the flexor reflex. To set the scene for these major achievements I first provide a brief account of the understanding of the spinal cord in "reflex" and "voluntary" motor activities from the beginning of the 20th century. Next, subsequent work is reviewed on the convergence on spinal interneurons from segmental sensory afferents and descending motor pathways, much of which was performed and inspired by Anders Lundberg's group in Gothenburg, SWE. This work was the keystone for new hypotheses on the role of spinal circuits in normal motor control. Such hypotheses were later tested under more natural conditions; either by recording directly from interneurons in reduced animal preparations or by use of indirect non-invasive techniques in humans performing normal movements. Some of this latter work is also reviewed. These developments would not have been possible without the preceding work on spinal reflexes by Eccles and Lundberg. Finally, there is discussion of how Eccles' work on spinal reflexes remains central (1) as new techniques are introduced on direct recording from interneurons in behaving animals; (2) in experiments on plastic neuronal changes in relation to motor learning and neurorehabilitation; (3) in experiments on transgenic animals uncovering aspects of human pathophysiology; and (4) in evaluating the function of genetically identified classes of neurons in studies on the development of the spinal cord.