Positive Feedback in a Brainstem Tactile Sensorimotor Loop

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Positive Feedback in a Brainstem Tactile Sensorimotor Loop Quoc-Thang Nguyen, David Kleinfeld  Neuron  Volume 45, Issue 3, Pages 447-457 (February 2005) DOI: 10.1016/j.neuron.2004.12.042 Copyright © 2005 Elsevier Inc. Terms and Conditions

Figure 1 The Vibrissa Trigeminal Loop and Basic Response (A) The vibrissa trigeminal loop in the rat brainstem. The gray dashed line indicates supratrigeminal sensorimotor loops. The black dashed lines outline the boundaries of the slice preparation. BL, buccolabialis branch of facial nerve; FN, facial nucleus; IoN, infraorbital branch of trigeminal nerve; PrV, principalis nucleus; RF, pontomedullary reticular formation; SpVc, spinal trigeminal nucleus pars caudalis; SpVi, spinal trigeminal nucleus pars interpolaris; TG, trigeminal ganglion; TN, trigeminal nuclear complex. (B) Brainstem slice from a P12 rat. The dashed box labeled FN outlines vibrissa motoneurons retrogradely labeled by Evans Blue. The dashed box labeled TN surrounds the trigeminal nuclear sensory complex. IO, inferior olive; V, trigeminal nerve. (C) EPSPs in vibrissa motoneurons following stimulation of the ipsilateral IoN in a slice from a P9 animal. Upper trace, response obtained with a 10 V IoN stimulation; lower trace, control with subthreshold 0.5 V stimulation; inset, average of 20 responses at 10 V stimulation. (D) EPSPs following stimulation of the ipsilateral trigeminal nerve in a slice from a P12 animal. Upper trace, response obtained with suprathreshold stimulation for spikes; lower trace, control with 2 V stimulation; inset, average of 20 responses with increasing stimulation voltage. (E) Trial-to-trial variability of EPSPs following stimulation of the ipsilateral trigeminal nerve in a slice from a P12 animal. (F) Postsynaptic depression of the motoneuron response. Traces are averages of ten trials. Neuron 2005 45, 447-457DOI: (10.1016/j.neuron.2004.12.042) Copyright © 2005 Elsevier Inc. Terms and Conditions

Figure 2 Nature of the Motoneuron EPSPs (A) Averaged EPSPs (20 trials) at different membrane potentials. (B) Distribution of the latency to the first EPSP, Δt as defined in (A) (white bars, cluster denoted Δt1; gray bars, cluster denoted Δt2) in P7 to P9 (hatched) and P12 to P15 preparations (clear). (C) Distribution of the maximal amplitude, A (see [A]) versus EPSP latency in vibrissa motoneurons from P12 to P15 animals. (D) Trial and preparation averaged intracellular response to trigeminal nerve stimulation. The responses from ten trials each of 25 motoneurons from P10 to P15 animals were averaged together; mean, black trace; SEM, gray band. Neuron 2005 45, 447-457DOI: (10.1016/j.neuron.2004.12.042) Copyright © 2005 Elsevier Inc. Terms and Conditions

Figure 3 EMG Responses Elicited by Stimulation of the IoN in Ketamine-Xylazine Anesthetized Rats (A) Experimental setup. (B) Response latencies Δt1 and Δt2 in a single trial. (C) Simultaneous recordings of EMG signals in ipsilateral intrinsic and extrinsic vibrissa muscles in three successive trials. The IoN was stimulated with a 5 mA, 200 μs current pulse delivered every 2 s. Average EMGs recorded before (light gray) and after (dark gray) transection of the buccolabialis branch are shown for a different rat. Traces are averages of 10 consecutive responses following IoN stimulation at 10 Hz with 8 mA, 100 μs pulses. (D) Distribution of response latencies in four rats (50 trials per rat). (E) Integral of rectified EMG responses of intrinsic muscles (∫dt |EMG|, inset, gray area in bottom trace) in four rats in response to increase in ION stimulation intensity (100 μs duration, delivered every 2 s). Each symbol represents a different animal. Each data point is the average (±SEM) of 10 to 20 trials; the line shows the trend. (F) Frequency-dependent depression of the averaged rectified trigeminally-evoked EMG response in intrinsic muscles and in the levator labii extrinsic muscle of the same rat. Each trace is the average of 20 rectified EMG records in response to repetitive IoN stimulations with 3 mA, 100 μs pulses. Arrows indicate IoN stimulation, which produces a stimulus artifact. Notice that the EMG responses consist of mixed Δt1 and Δt2 components in the intrinsic muscles and only a Δt1 component in the extrinsic muscle. Neuron 2005 45, 447-457DOI: (10.1016/j.neuron.2004.12.042) Copyright © 2005 Elsevier Inc. Terms and Conditions

Figure 4 EMG Responses Elicited by Artificial Whisking in Ketamine-Xylazine Anesthetized Rats (A) Experimental setup. (B) Response latencies Δt1 and Δt2 in a single trial. (C) Simultaneous recordings of EMG signals in ipsilateral intrinsic and extrinsic vibrissa muscles in three successive trials. The buccolabialis branch was stimulated with a 3 mA, 50 μs pulse delivered every 2 s. Average EMGs recorded before (light gray) and after (dark gray) transection of the infraorbital branch are shown for another rat. Traces are averages of 40 consecutive responses following buccolabialis stimulation at 0.5 Hz with 6 mA, 50 μs pulses. (D) Distribution of response latencies in four rats (50 trials per rat). Neuron 2005 45, 447-457DOI: (10.1016/j.neuron.2004.12.042) Copyright © 2005 Elsevier Inc. Terms and Conditions

Figure 5 EMG Responses Elicited by Passive Vibrissa Deflection in Ketamine-Xylazine Anesthetized Rats (A) Experimental setup for mechanical stimulation of the vibrissae. (B) EMG responses in intrinsic vibrissa muscles and in the levator labii superioris extrinsic muscle in response to a 20 Hz train of protractions in three consecutive trials. (C) Distribution of EMG response latencies in intrinsic muscles upon passive protraction (black bars) or simulated whisking through rhythmic stimulation of the buccolabialis branch of the facial nerve (gray bars) in five rats (100–116 trials per rat). Neuron 2005 45, 447-457DOI: (10.1016/j.neuron.2004.12.042) Copyright © 2005 Elsevier Inc. Terms and Conditions

Figure 6 EMG Responses Elicited by Vibrissa Contact in Ketamine-Xylazine Anesthetized Rats (A) Experimental setup for contact experiment during simulated whisking. (B) Contact signal (left traces) and consecutive averaged rectified EMG responses recorded simultaneously in intrinsic (middle traces) and extrinsic (right traces) vibrissa muscles. Each trace is the average of 30 sweeps, obtained by stimulating the buccolabialis branch of the facial nerve at 8 Hz with 9 mA, 50 μs pulses. In the <|EMGintrinsic|> records, the initial compound muscle action potential (e.g., Figure 4B) was truncated for the sake of clarity. (C) Distribution of the integral of the rectified EMG response in intrinsic (left graph) and levator labii superioris extrinsic (right graph) muscles during simulated whisking with (gray bars) or without contact (black bars) in one rat. The inset in each graph is a plot of the cumulated probability density function of the integral of the rectified EMG response in the presence or the absence of contact. (D) Distribution of the integral of the rectified EMG response in intrinsic (left graph; n = 7 rats) and extrinsic (right graph; n = 6 rats) muscles during simulated whisking with (gray bars) or without contact (black bars). Neuron 2005 45, 447-457DOI: (10.1016/j.neuron.2004.12.042) Copyright © 2005 Elsevier Inc. Terms and Conditions