Why are threatening experiences remembered so well? Insights into memory strengthening from protocols of gradual aversive learning

Aversive experiences often result in strong and persistent memory traces, which can sometimes lead to conditions such as Post-Traumatic Stress Disorder or phobias. Aversive stimulation tests are key tools in psychology and neuroscience for studying learning and memory. These tests typically use electric shocks as the unconditioned stimulus, allowing for precise control over the aversive content of the learning event. This feature has led to extensive research applying these tests with varying shock intensities to examine differences in learning, behavior, and memory formation between low- and high-aversive experiences. This line of research is particularly valuable for understanding the neurobiology underlying memory strengthening, but, to our knowledge, no review has yet compiled and organized the findings from this specific methodology. In this comprehensive review, we focus primarily on animal studies that have employed the same aversive test (i.e. Fear Conditioning, Passive Avoidance, Active Avoidance or Operant boxes) at different intensities. We will first outline and briefly describe the main aversive learning paradigms used in this field. Next, we will examine the relationship between aversiveness and memory strength. Finally, we will explore the neurobiological insights these studies have revealed over the years. Our aim is to gain a better understanding of how the nervous system gradually strengthens memory, while also addressing the remaining gaps and challenges in this area of research.