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Furthermore, the ability to observe spatial changes in biosensor signals, either through direct spatial quantification or the use of subcellular targeting, has not only confirmed the existence of spatially compartmentalized signaling domains but also provided the means with which to probe the mechanisms that give rise to this intricate spatial organization (Section 4).While the multifaceted nature of signal transduction can make it difficult to directly infer the critical regulators that underlie the observations made using fluorescent biosensors, computational models have been a useful companion to biosensor-based studies, as computational models benefit from the high temporal and spatial resolution of fluorescent biosensors and can in turn can quantitatively evaluate potential hypotheses and probe aspects of biology are not directly measurable experimentally (Section 5)