Phasic Firing Neurons Research Articles

Intrinsic properties of avian interaural level difference sound localizing neurons

Intrinsic properties of neurons are one major determinant for how neurons respond to their synaptic inputs and shape their outputs in neural circuits. Here, we studied the intrinsic properties of neurons in the chicken posterior portion of the dorsal nucleus of the lateral lemniscus (LLDp), the first interaural level difference (ILD) encoder of the avian auditory pathway. Using whole-cell recordings in brain slices, we revealed that the LLDp is composed of a heterogeneous neuron population based on their firing patterns. LLDp neurons were broadly classified as either phasic or tonic firing neurons, with further classification applied to tonic firing neurons, such as regular (most dominant, n = 82 out of 125 cells, 65.6%), pauser, or adaptive firing. Neurons with different firing patterns were distributed about evenly across the dorsoventral as well as mediolateral axis of LLDp. Phasic firing neurons were of faster membrane time constant, and lower excitability than tonic firing neurons. The action potentials (APs) elicited at the current thresholds displayed significant differences in first spike latency, AP peak amplitude, half-width, and maximal rising and falling rates. Interestingly, for APs elicited at suprathreshold currents (400 pA above thresholds), some of the differences diminished while a few others emerged. Remarkably, most parameters of the APs at thresholds were significantly different from those of APs at suprathresholds. Combined with our previous study (Curry and Lu, 2016), the results lend support to the two-cell type model for ILD coding in the avian system.

Phasic and tonic cell types in the zebra finch auditory caudal mesopallium.

The caudal mesopallium (CM) is a cortical-level area in the songbird auditory pathway where selective, invariant responses to familiar songs emerge. To characterize the cell types that perform this computation, we made whole cell recordings from brain slices in juvenile zebra finches ( Taeniopygia guttata) of both sexes. We found three groups of putatively excitatory neurons with distinct firing patterns. Tonic cells produced sustained responses to depolarizing step currents, phasic cells produced only a few spikes at the onset, and an intermediate group was also phasic but responded for up to a few hundred milliseconds. Phasic cells had smaller dendritic fields, higher resting potentials, and strong low-threshold outward rectification. Pharmacological treatment with voltage-gated potassium channel antagonists 4-aminopyridine and α-dendrotoxin converted phasic to tonic firing. When stimulated with broadband currents, phasic cells fired coherently with frequencies up to 20-30 Hz, whereas tonic neurons were more responsive to frequencies around 0-10 Hz. The distribution of peak coherence frequencies was similar to the distribution of temporal modulation rates in zebra finch song. We reproduced these observations in a single-compartment biophysical model by varying cell size and the magnitude of a slowly inactivating, low-threshold potassium current ( ILT). These data suggest that intrinsic dynamics in CM are matched to the temporal statistics of conspecific song. NEW & NOTEWORTHY In songbirds, the caudal mesopallium is a key brain area involved in recognizing the songs of other individuals. This study identifies three cell types in this area with distinct firing patterns (tonic, phasic, and intermediate) that reflect differences in cell size and a low-threshold potassium current. The phasic-firing neurons, which do not have a counterpart in mammalian auditory cortex, are better able to follow rapid modulations at the frequencies found in song.

Slope-Based Stochastic Resonance: How Noise Enables Phasic Neurons to Encode Slow Signals

Fundamental properties of phasic firing neurons are usually characterized in a noise-free condition. In the absence of noise, phasic neurons exhibit Class 3 excitability, which is a lack of repetitive firing to steady current injections. For time-varying inputs, phasic neurons are band-pass filters or slope detectors, because they do not respond to inputs containing exclusively low frequencies or shallow slopes. However, we show that in noisy conditions, response properties of phasic neuron models are distinctly altered. Noise enables a phasic model to encode low-frequency inputs that are outside of the response range of the associated deterministic model. Interestingly, this seemingly stochastic-resonance (SR) like effect differs significantly from the classical SR behavior of spiking systems in both the signal-to-noise ratio and the temporal response pattern. Instead of being most sensitive to the peak of a subthreshold signal, as is typical in a classical SR system, phasic models are most sensitive to the signal's rising and falling phases where the slopes are steep. This finding is consistent with the fact that there is not an absolute input threshold in terms of amplitude; rather, a response threshold is more properly defined as a stimulus slope/frequency. We call the encoding of low-frequency signals with noise by phasic models a slope-based SR, because noise can lower or diminish the slope threshold for ramp stimuli. We demonstrate here similar behaviors in three mechanistic models with Class 3 excitability in the presence of slow-varying noise and we suggest that the slope-based SR is a fundamental behavior associated with general phasic properties rather than with a particular biological mechanism.

Task-dependent encoding of space and events by striatal neurons is dependent on neural subtype

The dorsal striatum plays a critical role in procedural learning and memory. Current models of basal ganglia assume that striatal neurons and circuitry are critical for the execution of overlearned, habitual sequences of action. However, less is known about how the striatum encodes task information that guides the performance of actions in procedural tasks. To explore the striatal encoding of task information, we compared the behavioral correlates of striatal neurons tested in two tasks: a multiple T-maze task in which reward delivery was entirely predictable based on spatial cues (the Multiple-T task), and a task in which rats ran on a rectangular track, but food delivery depended on the distance traveled on the track and was not dependent solely on spatial location (the Take-5 task). Striatal cells recorded on these tasks were divisible into three cell types: phasic-firing neurons (PFNs), tonically firing neurons (TFNs), and high-firing neurons (HFNs) and similar proportions of each cell type were found in each task. However, the behavioral correlates of each cell type were differentially sensitive to the type of task rats were performing. PFNs were responsive to specific task-parameters on each task. TFNs showed reliable burst-and-pause responses following food delivery and other events that were consistent with tonically active neurons (TANs) on the Take-5 (non-spatial) task but not on the Multiple-T (spatial) task. HFNs showed spatial oscillations on the Multiple-T (spatial) task but not the Take-5 (non-spatial) task. Reconstruction of the rats' position on the maze was highly accurate when using striatal ensembles recorded on the Multiple-T (spatial) task, but not when using ensembles recorded on the Take-5 (non-spatial) task. In contrast, reconstruction of time following food delivery was successful in both tasks. The results indicated a strong task dependency of the quality of the spatial, but not the reward-related, striatal representations on these tasks. These results suggest that striatal spatial representations depend on the degree to which spatial task-parameters can be unambiguously associated with goals.

Subpopulation of Dorsal Horn Neurons Displays Enhanced N -methyl-d-aspartate Receptor Function after Chronic Morphine Exposure

Morphine tolerance may be attributed to enhancement of glutamatergic neurotransmission, in particular to increased function of the N-methyl-D-aspartate (NMDA) receptor. The cellular mechanisms responsible for these changes remain poorly defined. The authors identified and characterized a specific subpopulation of dorsal horn neurons, displaying NMDA receptor plasticity in response to chronic morphine administration. The authors undertook current clamped and voltage clamped recordings of NMDA receptor-mediated responses from cultured rat dorsal horn neurons that were untreated or treated for 7 days with 1 or 100 microm morphine. Smaller (capacitance < or = 22 pF), tonic firing neurons showed a significantly enhanced NMDA receptor-mediated peak current after prolonged morphine treatment, whereas larger and phasic firing neurons showed no enhancement. With high-concentration but not low-concentration morphine treatment, Mg2+ blockade of NMDA receptors at resting membrane potentials was reduced. Furthermore, the chronic opioid-induced increase in NMDA current was attenuated by pretreatment with either a mu-opioid receptor inhibitor (naloxone) or an NMDA receptor inhibitor (2-amino-5-phosphonovalerate) (low-concentration > high-concentration morphine). In an electrophysiologically defined subpopulation of dorsal horn neurons, enhanced NMDA receptor function after chronic morphine exposure was shown to be mechanistically dependent on morphine concentration and sensitive to both NMDA and mu-opioid receptor antagonism. Therefore, these changes observed in this population of sensory spinal neurons can be used to study the development and prevention of opioid tolerance described in multiple laboratory and clinical reports.

Thermosensitivity of neurons in the paraventricular nucleus of the rat slice preparation

The thermosensitivity of 65 spontaneouslu active neurons in the paraventricular nucleus (PVN) was investigated by extracellular recording in the rat hypothalamic slice preparation. The firing rate of these cells was comparatively low, ranging from 0.03 to 10.0 (mean-2.46) impulses/s at 37 °C, and only a minority showed a phasic firing pattern. Of 65 neurons tested, 23 (35%) increased their firing rate when the slice was warmed (warm-sensitive neurons) and 9 (14%) showed the opposite response (cold-sensitive neurons). Thermosensitivity was also tested in solutions with reduced [Ca 2+] and high [Mg 2+]. Eight out of 10 warm-sensitive neurons and 5 of 7 cold-sensitive neurons retained thermosensitivity after synaptic blockade. Out of 6 phasic firing neurons tested, one showed warm-sensitivity and another one showed cold-sensitivity. The thermosensitive neurons were diffusely distributed throughout the PVN and were not located in particular areas of the nucleus. Thus a group of cells in the PVN, including probably both magno- and parvocellular neurons, showed an inherent thermosensitivity, which suggests an important role for the PVN in thermoregulation.

Angiotensin II sensitive neurons in the supraoptic nucleus, subfornical organ and anteroventral third ventricle of rats in vitro

The angiotensin II (AII) sensitivity of neurons in the supraoptic nucleus (SON), subfornical organ (SFO) and the region near the anteroventral part of the third ventricle (AV3V) was investigated using extracellular recording in the rat brain slice preparation by adding AII (10 −10–10 −6 M) to the perfusion medium. Forty seven (44%) of 106 SON neurons, 62 (66%) of 94 SFO neurons and 28 (33%) of 86 AV3V neurons were excited by AII. One cell was inhibited by AII in the SON and one in the SFO. The threshold concentration to evoke responses in the SON neurons was approximately 10 −9 M, but nuerons in the SFO and AV3V showed clear excitatory responses to AII at 10 −10 M. In the SON, 18 (40%) of 45 phasic firing neurons (putative vasopressin neurons) and 29 (48%) of 61 nonphasic firing neurons (including putative oxytocin neurons) were excited by AII. The excitatory effect of AII was reversibly antagonized by a specific antagonist saralasin and persisted after synaptic blockade in medium with low [Ca 2+] and high [Mg 2+]. We conclude that AII can stimulate both vasopressin and oxytocin release, acting directly upon SON neurons and also that both the SFO and AV3V are important receptive sites for AII (although the SFO is relatively more sensitive) which contributes SON input and modulates release of these hormones.