Do any nonhuman animals have hedonically valenced experiences not directly caused by stimuli in their current environment? Do they, like us humans, experience anticipated or previously experienced pains and pleasures as respectively painful and pleasurable? We review evidence from comparative neuroscience about hippocampus-dependent simulation in relation to this question. Hippocampal sharp-wave ripples and theta oscillations have been found to instantiate previous and anticipated experiences. These hippocampal activations coordinate with neural reward and fear centers as well as sensory and cortical areas in ways that are associated with conscious episodic mental imagery in humans. Moreover, such hippocampal "re- and preplay" has been found to contribute to instrumental decision making, the learning of value representations, and the delay of rewards in rats. The functional and structural features of hippocampal simulation are highly conserved across mammals. This evidence makes it reasonable to assume that internally triggered experiences of hedonic valence (IHVs) are pervasive across (at least) all mammals. This conclusion has important welfare implications. Most prominently, IHVs act as a kind of "welfare multiplier" through which the welfare impacts of any given experience of pain or pleasure are increased through each future retrieval. However, IHVs also have practical implications for welfare assessment and cause prioritization.
Keywords: affect; animal welfare; comparative psychology; emotion; episodic memory; hedonic valence; hippocampal re- and preplay; mental time travel.