Complex dendrites in general present formidable challenges to understanding neuronal information processing. To circumvent the difficulty, a prevalent viewpoint simplifies the neuronal morphology as a point representing the soma, and the excitatory and inhibitory synaptic currents originated from the dendrites are treated as linearly summed at the soma. Despite its extensive applications, the validity of the synaptic current description remains unclear, and the existing point neuron framework fails to characterize the spatiotemporal aspects of dendritic integration supporting specific computations. Using electrophysiological experiments, realistic neuronal simulations, and theoretical analyses, we demonstrate that the traditional assumption of linear summation of synaptic currents is oversimplified and underestimates the inhibition effect. We then derive a form of synaptic integration current within the point neuron framework to capture dendritic effects. In the derived form, the interaction between each pair of synaptic inputs on the dendrites can be reliably parameterized by a single coefficient, suggesting the inherent low-dimensional structure of dendritic integration. We further generalize the form of synaptic integration current to capture the spatiotemporal interactions among multiple synaptic inputs and show that a point neuron model with the synaptic integration current incorporated possesses the computational ability of a spatial neuron with dendrites, including direction selectivity, coincidence detection, logical operation, and a bilinear dendritic integration rule discovered in experiment. Our work amends the modeling of synaptic inputs and improves the computational power of a modeling neuron within the point neuron framework.
Neurons process information via integration of synaptic inputs from dendrites. Many experimental results demonstrate dendritic integration could be highly nonlinear, yet few theoretical analyses have been performed to obtain a precise quantitative characterization analytically. Based on asymptotic analysis of a two-compartment passive cable model, given a pair of time-dependent synaptic conductance inputs, we derive a bilinear spatiotemporal dendritic integration rule. The summed somatic potential can be well approximated by the linear summation of the two postsynaptic potentials elicited separately, plus a third additional bilinear term proportional to their product with a proportionality coefficient . The rule is valid for a pair of synaptic inputs of all types, including excitation-inhibition, excitation-excitation, and inhibition-inhibition. In addition, the rule is valid during the whole dendritic integration process for a pair of synaptic inputs with arbitrary input time differences and input locations. The coefficient is demonstrated to be nearly independent of the input strengths but is dependent on input times and input locations. This rule is then verified through simulation of a realistic pyramidal neuron model and in electrophysiological experiments of rat hippocampal CA1 neurons. The rule is further generalized to describe the spatiotemporal dendritic integration of multiple excitatory and inhibitory synaptic inputs. The integration of multiple inputs can be decomposed into the sum of all possible pairwise integration, where each paired integration obeys the bilinear rule. This decomposition leads to a graph representation of dendritic integration, which can be viewed as functionally sparse.
The Granger causality (GC) analysis has been extensively applied to infer causal interactions in dynamical systems arising from economy and finance, physics, bioinformatics, neuroscience, social science, and many other fields. In the presence of potential nonlinearity in these systems, the validity of the GC analysis in general is questionable. To illustrate this, here we first construct minimal nonlinear systems and show that the GC analysis fails to infer causal relations in these systems-it gives rise to all types of incorrect causal directions. In contrast, we show that the time-delayed mutual information (TDMI) analysis is able to successfully identify the direction of interactions underlying these nonlinear systems. We then apply both methods to neuroscience data collected from experiments and demonstrate that the TDMI analysis but not the GC analysis can identify the direction of interactions among neuronal signals. Our work exemplifies inference hazards in the GC analysis in nonlinear systems and suggests that the TDMI analysis can be an appropriate tool in such a case.
It has been discovered recently in experiments that the dendritic integration of excitatory glutamatergic inputs and inhibitory GABAergic inputs in hippocampus CA1 pyramidal neurons obeys a simple arithmetic rule as , where , and are the respective voltage values of the summed somatic potential, the excitatory postsynaptic potential (EPSP) and the inhibitory postsynaptic potential measured at the time when the EPSP reaches its peak value. Moreover, the shunting coefficient in this rule only depends on the spatial location but not the amplitude of the excitatory or inhibitory input on the dendrite. In this work, we address the theoretical issue of how much the above dendritic integration rule can be accounted for using subthreshold membrane potential dynamics in the soma as characterized by the conductance-based integrate-and-fire (I&F) model. Then, we propose a simple I&F neuron model that incorporates the spatial dependence of the shunting coefficient by a phenomenological parametrization. Our analytical and numerical results show that this dendritic-integration-rule-based I&F (DIF) model is able to capture many experimental observations and it also yields predictions that can be used to verify the validity of the DIF model experimentally. In addition, the DIF model incorporates the dendritic integration effects dynamically and is applicable to more general situations than those in experiments in which excitatory and inhibitory inputs occur simultaneously in time. Finally, we generalize the DIF neuronal model to incorporate multiple inputs and obtain a similar dendritic integration rule that is consistent with the results obtained by using a realistic neuronal model with multiple compartments. This generalized DIF model can potentially be used to study network dynamics that may involve effects arising from dendritic integrations.
This paper presents a fully integrated reconfigurable all-band RF transceiver for GPS/GLONASS/ Galileo/Beidou in 55-nm CMOS. The transceiver incorporates three low-IF receivers (RXs) and one direct up-conversion BPSK transmitter (TX), which can be configured to receive any two global navigation satellite system (GNSS) signals or switched to process the Chinese Beidou(I) signals. A switching module is integrated to provide the connectivity between different RF front-end and IF channel (IFC), which will effectively simplify the design complexity of the IFC and save power consumption, while the GNSS signals are received. A flexible frequency plan with two frequency synthesizers is utilized to satisfy different local oscillator requirements of the transceiver. An optimized automatic frequency calibration scheme using an error compensation logic enables fast and high-precision calibration process for optimum phase-locked loop operation. Several digitally assisted calibration modules are integrated to ensure that the chip performance only shows a weak process, voltage and temperature (PVT) dependence. While drawing about 21.5-30.2 mA per RX channel from a 1.2-V supply, the RXs achieve an image rejection ratio more than 49 dB after I/Q mismatch calibration, an automatic gain control range of 88 dB, and an input-referred 1 dB compression point of better than −25 dBm with a minimum noise figure of about 2 dB. The output power of the TX is about 5 dBm with about 6% error vector magnitude (EVM) and 30-mA current from a 1.2-V supply. The whole transceiver consumes a die area of 2.8 × 3 mm 2 .
Significance In the neocortex, while early sensory areas encode and process external inputs rapidly, higher-association areas are endowed with slow dynamics suitable for accumulating information over time. Such a hierarchy of temporal response windows along the cortical hierarchy naturally emerges in a model of multiareal primate cortex. This finding raises the question of why diverse temporal modes are not mixed in roughly the same way across the whole cortex, despite high connection density and an abundance of feedback loops. We investigate this question by mathematically analyzing the anatomically based network model of macaque cortex and theoretically show that three sufficient conditions of synaptic excitation and inhibition give rise to timescale segregation in a hierarchy, a functionally important characteristic of the cortex.
We address the question of how a neuron integrates excitatory (E) and inhibitory (I) synaptic inputs from different dendritic sites. For an idealized neuron model with an unbranched dendritic cable, we construct its Green's function and carry out an asymptotic analysis to obtain its solutions. Using these asymptotic solutions, in the presence of E and I inputs, we can successfully reveal the underlying mechanisms of a dendritic integration rule, which was discovered in a recent experiment. Our analysis can be extended to the multi-branch case to characterize the E-I dendritic integration on any branches. The novel characterization is confirmed by the numerical simulation of a biologically realistic neuron.
Interneurons are important for computation in the brain, in particular, in the information processing involving the generation of theta oscillations in the hippocampus. Yet the functional role of interneurons in the theta generation remains to be elucidated. Here we use time-delayed mutual information to investigate information flow related to a special class of interneurons—theta-driving neurons in the hippocampal CA1 region of the mouse—to characterize the interactions between theta-driving neurons and theta oscillations. For freely behaving mice, our results show that information flows from the activity of theta-driving neurons to the theta wave, and the firing activity of theta-driving neurons shares a substantial amount of information with the theta wave regardless of behavioral states. Via realistic simulations of a CA1 pyramidal neuron, we further demonstrate that theta-driving neurons possess the characteristics of the cholecystokinin-expressing basket cells (CCK-BC). Our results suggest that it is important to take into account the role of CCK-BC in the generation and information processing of theta oscillations.
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