Measuring spectrally-resolved information transfer for sender- and receiver-specific frequencies

Information transfer, measured by transfer entropy, is a key component of distributed computation. It is therefore important to understand the pattern of information transfer in order to unravel the distributed computational algorithms of a system. Since in many natural systems distributed computation is thought to rely on rhythmic processes a frequency resolved measure of information transfer is highly desirable. Here, we present a novel algorithm, and its efficient implementation, to identify separately frequencies sending and receiving information in a network. Our approach relies on the invertible maximum overlap discrete wavelet transform (MODWT) for the creation of surrogate data in the computation of transfer entropy and entirely avoids filtering of the original signals. The approach thereby avoids well-known problems due to phase shifts or the ineffectiveness of filtering in the information theoretic setting. We also show that measuring frequency-resolved information transfer is a partial information decomposition problem that cannot be fully resolved to date and discuss the implications of this issue. Last, we evaluate the performance of our algorithm on simulated data and apply it to human magnetoencephalography (MEG) recordings and to local field potential recordings in the ferret. In human MEG we demonstrate top-down information flow in temporal cortex from very high frequencies (above 100Hz) to both similarly high frequencies and to frequencies around 20Hz, i.e. a complex spectral configuration of cortical information transmission that has not been described before. In the ferret we show that the prefrontal cortex sends information at low frequencies (4-8 Hz) to early visual cortex (V1), while V1 receives the information at high frequencies (> 125 Hz). Author Summary Systems in nature that perform computations typically consist of a large number of relatively simple but interacting parts. In human brains, for example, billions of neurons work together to enable our cognitive abilities. This well-orchestrated teamwork requires information to be exchanged very frequently. In many cases this exchange happens rhythmically and, therefore, it seems beneficial for our understanding of physical systems if we could link the information exchange to specific rhythms. We here present a method to determine which rhythms send, and which rhythms receive information. Since many rhythms can interact at both sender and receiver side, we show that the interpretation of results always needs to consider that the above problem is tightly linked to partial information decomposition - an intriguing problem from information theory only solved recently, and only partly. We applied our novel method to information transfer in the human inferior temporal cortex, a brain region relevant for object perception, and unexpectedly found information transfer originating at very high frequencies at 100Hz and then forking to be received at both similarly high but also much lower frequencies around 20Hz. These results overturn the current standard assumption that low frequencies send information to high frequencies.