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The Inhibitory Thermal Effects of Focused Ultrasound on an Identified, Single Motoneuron

Morgan N. Collins, Wynn Legon, Karen A. Mesce

eneuro 2021, 8, ENEURO.0514-20.2021 · 10.1523/eneuro.0514-20.2021

invertebratehealthyinvasive electrophysiology

Abstract

Focused ultrasound (US) is an emerging neuromodulation technology that has gained much attention because of its ability to modulate, noninvasively, neuronal activity in a variety of animals, including humans. However, there has been considerable debate about exactly which types of neurons can be influenced and what underlying mechanisms are in play. Are US-evoked motor changes driven indirectly by activated mechanosensory inputs, or more directly via central interneurons or motoneurons? Although it has been shown that US can mechanically depolarize mechanosensory neurons, there are no studies that have yet tested how identified motoneurons respond directly to US and what the underlying mechanism might be. Here, we examined the effects of US on a single, identified motoneuron within a well-studied and tractable invertebrate preparation, the medicinal leech, Hirudo verbana Our approach aimed to clarify single neuronal responses to US, which may be obscured in other studies whereby US is applied across a diverse population of cells. We found that US has the ability to inhibit tonic spiking activity through a predominately thermal mechanism. US-evoked effects persisted after blocking synaptic inputs, indicating that its actions were direct. Experiments also revealed that US-comparable heating blocked the axonal conduction of spontaneous action potentials. Finally, we found no evidence that US had significant mechanical effects on the neurons tested, a finding counter to prevailing views. We conclude that a non-sensory neuron can be directly inhibited via a thermal mechanism, a finding that holds promise for clinical neuromodulatory applications.

Abstract via europepmc.

Speciesmedicinal leech (Hirudo verbana)
Subjects10, 26, 21, 4swept preparations
Sessions per subject1
Randomisednot reported
Blindingnot reported
Sham / controlno treatment control
Auditory controlnot reported
Readout timingboth
Anaesthesianot reported
Readoutsinvasive electrophysiologyextracellular suction-electrode recording of DP nerve/DE-3 spiking; intracellular sharp-electrode recording of DE-3 soma
Direction of effectbidirectionalOf 18 responsive DE-3 motoneurons, 13 showed inhibition (mean 43.3% decrease in firing) and 4 showed excitation (mean 60.7% increase); the predominant, direct, thermally-mediated effect was inhibition via axonal conduction block.
Adverse eventsobservedA minority of strongly inhibited nerves failed to recover baseline firing for the remainder of viability (e.g., one nerve fired only once 60 s after stimulus end); most nerves (77.8%) recovered within 20% of baseline.

Exposures

Exposure 1: 960-kHz pulsed US applied to the dorsal posterior (DP) nerve / DE-3 motoneuron axon

Target: leech neurons — “DE-3 motoneuron axon in the dorsal posterior (DP) nerve of the medicinal leech
Device: Sonic Concepts · Sonic Concepts · H-102MR

Pulse timing
Waveformpulsed
Fundamental frequency (kHz)960✓✓
Pulse duration (ms)0.3✓✓
Pulse repetition frequency (Hz)1,000✓✓
Duty cycle (%)30pulse duration × PRF gives 30%✓✓
Sonication duration (s)0.1, 0.316, 1, 3.16, 10, 30swept✓✓
Pressure and intensity, by domain
Free-field pressure (kPa)660, 1,300swept✓✓
Free-field Isppa (W/cm²)14.52✓✓
Free-field Ispta (W/cm²)4.84✓✓
In-situ estimatenot applicable
In-situ pressure (kPa)not applicable
In-situ Isppa (W/cm²)not applicable
In-situ Ispta (W/cm²)not applicable
Protocol, in the paper’s words

US pulses of 290 cycles (~300 us) applied at a 1-kHz pulse repetition frequency (intrapulse duty cycle 30%) for a 30-s stimulus train, within a 90-s trial (30 s baseline, 30 s stimulus, 30 s recovery, with longer recovery for shorter-duration pilot trials). Five shorter application durations (100 ms-10 s) were piloted first and failed to reliably modulate firing before 30 s was adopted as the standard duration.

Flags from extraction

  • exposures[0].timing.pulse_duration_msFigure 2 legend states 'Each US pulse was 300 ms in duration', but the Methods text states pulses were '290 cycles and were 300 <micro>s [us] in duration' (consistent with 290 cycles at 960 kHz ~=302 us); the figure legend's 'ms' appears to be a units/typo error, so 300 us (0.3 ms) is used.
  • exposures[0].timing.duty_cycle_pctFigure 2 legend gives two different duty-cycle figures in the same sentence: '50% duty cycle' and 'intrapulse duty cycle of 30%'. 30% is consistent with the stated 300 us pulse duration and 1 kHz PRF (1 ms period); 50% is not explained and may refer to a different (unspecified) duty parameter.
  • exposures[0].timing.pulse_repetition_frequency_hzMethods text states 'We applied 500 pulses/s at a 1-kHz pulse repetition frequency', which is internally inconsistent (500 pulses/s implies 500 Hz, not 1 kHz); 1 kHz is used here as the explicitly labeled 'pulse repetition frequency'.
  • exposures[0].unspecified_domain.pressure_kpaDomain (free-field vs in-situ) is not stated by the paper; the leech nerve was immersed directly in a saline bath (no intervening skull/tissue), and pressure was characterized by hydrophone in that setting.
  • n_subjects26 nerves were exposed to the main 30-s/960-kHz protocol (22 analysed after exclusions); additional, overlapping/non-overlapping subsets of nerves were also tested under modified conditions (dual-recording N=4, Ca2+-free saline N=10, low-heat latex-dish N=21, pressure-doubled N=4), which are not summed into a single total.