Explanatory models in neuroscience: Part 2 -- constraint-based intelligibility

Computational modeling plays an increasingly important role in neuroscience, highlighting the philosophical question of how computational models explain. In the context of neural network models for neuroscience, concerns have been raised about model intelligibility, and how they relate (if at all) to what is found in the brain. We claim that what makes a system intelligible is an understanding of the dependencies between its behavior and the factors that are causally responsible for that behavior. In biological systems, many of these dependencies are naturally "top-down": ethological imperatives interact with evolutionary and developmental constraints under natural selection. We describe how the optimization techniques used to construct NN models capture some key aspects of these dependencies, and thus help explain why brain systems are as they are -- because when a challenging ecologically-relevant goal is shared by a NN and the brain, it places tight constraints on the possible mechanisms exhibited in both kinds of systems. By combining two familiar modes of explanation -- one based on bottom-up mechanism (whose relation to neural network models we address in a companion paper) and the other on top-down constraints, these models illuminate brain function.

Explanatory models in neuroscience: Part 1 -- taking mechanistic abstraction seriously

Despite the recent success of neural network models in mimicking animal performance on visual perceptual tasks, critics worry that these models fail to illuminate brain function. We take it that a central approach to explanation in systems neuroscience is that of mechanistic modeling, where understanding the system is taken to require fleshing out the parts, organization, and activities of a system, and how those give rise to behaviors of interest. However, it remains somewhat controversial what it means for a model to describe a mechanism, and whether neural network models qualify as explanatory. We argue that certain kinds of neural network models are actually good examples of mechanistic models, when the right notion of mechanistic mapping is deployed. Building on existing work on model-to-mechanism mapping (3M), we describe criteria delineating such a notion, which we call 3M++. These criteria require us, first, to identify a level of description that is both abstract but detailed enough to be "runnable", and then, to construct model-to-brain mappings using the same principles as those employed for brain-to-brain mapping across individuals. Perhaps surprisingly, the abstractions required are those already in use in experimental neuroscience, and are of the kind deployed in the construction of more familiar computational models, just as the principles of inter-brain mappings are very much in the spirit of those already employed in the collection and analysis of data across animals. In a companion paper, we address the relationship between optimization and intelligibility, in the context of functional evolutionary explanations. Taken together, mechanistic interpretations of computational models and the dependencies between form and function illuminated by optimization processes can help us to understand why brain systems are built they way they are.