The behavior of insects transcends elementary forms of adaptive responding to environmental changes. We discuss examples of exploration, instrumental and observational learning, expectation, learning in a social context, and planning of future actions. We show that learning about sensory cues allows insects to transfer flexibly their responses to novel stimuli attaining thereby different levels of complexity, from basic generalization to categorization and concept learning consistent with rule extraction. We argue that updating of existing memories requires multiple forms of memory processing. A key element in these processes is working memory, an active form of memory considered to allow evaluation of actions on the basis of expected outcome. We discuss which of these cognitive faculties can be traced to specific neural processes and how they relate to the overall organization of the insect brain.
What is the engram, the memory trace that stores the content of memory? Here, I argue that the engram is more than just the sum of all learning-related neural changes. Rather, it is an integrated part of the whole nervous system, from sensory integration to interneuron processing and motor control. Emphasis is given to those neural processing components that do not express themselves in the behavior of the animal. To date, the search for the olfactory engram in the bee brain has focused on network properties in the antennal lobe (AL), the input and output regions of the mushroom body region (calyx), the mushroom body extrinsic neurons, and the reward pathway. I conclude that the olfactory memory trace in the AL relates predominantly to attention-generating properties. The memory trace in the calyx is characterized by the high-order combinatorial integration of multiple sensory inputs. Mushroom body extrinsic neurons are tentatively related to multiple processing categories that represent the acquired values (e.g., favorable conditions and to be avoided condition) and provide neural commands for goal-directed behavior and decision making. The system of VUM neurons stores appetitive memory related to appetitive internal states and controls nonassociative and associative memories. A model is presented that aims to capture the multifaceted and distributed nature of the engram and may help to guide our future search of the engram at a systems level.
What is the engram, the memory trace that stores the content of memory? Here, I argue that the engram is more than just the sum of all learning-related neural changes. Rather, it is an integrated part of the whole nervous system, from sensory integration to interneuron processing and motor control. Emphasis is given to those neural processing components that do not express themselves in the behavior of the animal. To date, the search for the olfactory engram in the bee brain has focused on network properties in the antennal lobe (AL), the input and output regions of the mushroom body region (calyx), the mushroom body extrinsic neurons, and the reward pathway. I conclude that the olfactory memory trace in the AL relates predominantly to attention-generating properties. The memory trace in the calyx is characterized by the high-order combinatorial integration of multiple sensory inputs. Mushroom body extrinsic neurons are tentatively related to multiple processing categories that represent the acquired values (e.g., favorable conditions and to be avoided condition) and provide neural commands for goal-directed behavior and decision making. The system of VUM neurons stores appetitive memory related to appetitive internal states and controls nonassociative and associative memories. A model is presented that aims to capture the multifaceted and distributed nature of the engram and may help to guide our future search of the engram at a systems level.
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