RESUMEN
Understanding the neural bases of subjective experience remains one of the great challenges of the natural sciences. Higher-order theories of consciousness are typically defended by assessments of neural activity in higher cortical regions during perception, often with disregard to the nature of the neural computations that these regions execute. We have sought to refocus the problem toward identification of those neural computations that are necessary for subjective experience with the goal of defining the sorts of neural architectures that can perform these operations. This approach removes reliance on behaviour and brain homologies for appraising whether non-human animals have the potential to subjectively experience sensory stimuli. Using two basic principles-first, subjective experience is dependent on complex processing executing specific neural functions and second, the structure-determines-function principle-we have reasoned that subjective experience requires a neural architecture consisting of stacked forward models that predict the output of neural processing from inputs. Given that forward models are dependent on appropriately connected processing modules that generate prediction, error detection and feedback control, we define a minimal neural architecture that is necessary (but not sufficient) for subjective experience. We refer to this framework as the hierarchical forward models algorithm. Accordingly, we postulate that any animal lacking this neural architecture will be incapable of subjective experience.
RESUMEN
How subjective experience is realized in nervous systems remains one of the great challenges in the natural sciences. An answer to this question should resolve debate about which animals are capable of subjective experience. We contend that subjective experience of sensory stimuli is dependent on the brain's awareness of its internal neural processing of these stimuli. This premise is supported by empirical evidence demonstrating that disruption to either processing streams or awareness states perturb subjective experience. Given that the brain must predict the nature of sensory stimuli, we reason that conscious awareness is itself dependent on predictions generated by hierarchically organized forward models of the organism's internal sensory processing. The operation of these forward models requires a specialized neural architecture and hence any nervous system lacking this architecture is unable to subjectively experience sensory stimuli. This approach removes difficulties associated with extrapolations from behavioral and brain homologies typically employed in addressing whether an animal can feel. Using nociception as a model sensation, we show here that the Drosophila brain lacks the required internal neural connectivity to implement the computations required of hierarchical forward models. Consequently, we conclude that Drosophila, and those insects with similar neuroanatomy, do not subjectively experience noxious stimuli and therefore cannot feel pain.