During the process of skill learning, synaptic connections in our brains are modified to form motor memories of learned sensorimotor acts. The more plastic the adult brain is, the easier it is to learn new skills or adapt to neurological injury. However, if the brain is too plastic and the pattern of synaptic connectivity is constantly changing, new memories will overwrite old memories, and learning becomes unstable. This trade-off is known as the stability-plasticity dilemma. Here a theory of sensorimotor learning and memory is developed whereby synaptic strengths are perpetually fluctuating without causing instability in motor memory recall, as long as the underlying neural networks are sufficiently noisy and massively redundant. The theory implies two distinct stages of learning-preasymptotic and postasymptotic-because once the error drops to a level comparable to that of the noiseinduced error, further error reduction requires altered network dynamics. A key behavioral prediction derived from this analysis is tested in a visuomotor adaptation experiment, and the resultant learning curves are modeled with a nonstationary neural network. Next, the theory is used to model two-photon microscopy data that show, in animals, high rates of dendritic spine turnover, even in the absence of overt behavioral learning. Finally, the theory predicts enhanced task selectivity in the responses of individual motor cortical neurons as the level of task expertise increases. From these considerations, a unique interpretation of sensorimotor memory is proposed-memories are defined not by fixed patterns of synaptic weights but, rather, by nonstationary synaptic patterns that fluctuate coherently.hyperplastic | neural tuning S ensorimotor skill learning, like other types of learning, occurs through the general mechanism of experience-dependent synaptic plasticity (1, 2). As we learn a new skill (such as a tennis stroke) through extensive practice, synapses in our brain are modified to form a lasting motor memory of that skill. However, if synapses are overly pliable and in a state of perpetual flux, memories may not stabilize properly as new learning can overwrite previous learning. Thus, for any distributed learning system, there is inherent tension between the competing requirements of stability and plasticity (3): Synapses must be sufficiently plastic to support the formation of new memories, while changing in a manner that preserves the traces of old memories. The specific learning mechanisms by which these contradictory constraints are simultaneously fulfilled are one of neuroscience's great mysteries.The inescapability of the stability-plasticity dilemma, as faced by any distributed learning system, is shown in the cartoon neural network in Fig. 1A. Suppose that the input pattern of [0.6, 0.4] must be transformed into the activation pattern [0.5, 0.7] at the output layer. Given the initial connectivity of the network, the input transforms to the incorrect output [0.8, 0.2]. Through practice and a learning mechanism, the weig...