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Focused ultrasound transiently increases membrane conductance in isolated crayfish axon

Jen-Wei Lin, Feiyuan Yu, Wolfgang S. Müller, Gösta Ehnholm, Yoshio Okada

Journal of Neurophysiology 2019, 121, 480-489 · 10.1152/jn.00541.2018

ex vivo tissuehealthyinvasive electrophysiology

Abstract

We report a novel phenomenon produced by focused ultrasound (US) that may be important for understanding its effects on cell membranes. When a US burst (2.1 MHz, 1-mm focal diameter, 0.1-1 MPa) was focused on a motor axon of the crayfish neuromuscular junction, it consistently produced a fast hyperpolarization, which was followed or superseded by subthreshold depolarizations or action potentials in a stochastic manner. The depolarization persisted in the presence of voltage-gated channel blockers [1 µM TTX ( I Na ), 50 µM ZD7288 ( I h ), and 200 µM 4-aminopyridine ( I K )] and typically started shortly after the onset of a 5-ms US burst, with a mean latency of 3.35 ± 0.53 ms (SE). The duration and amplitude of depolarizations averaged 2.13 ± 0.87 s and 10.1 ± 2.09 mV, with a maximum of 200 s and 60 mV, respectively. The US-induced depolarization was always associated with a decrease in membrane resistance. By measuring membrane potential and resistance during the US-induced depolarization, the reversal potential of US-induced conductance ( g us ) was estimated to be -8.4 ± 2.3 mV, suggesting a nonselective conductance. The increase in g us was 10-100 times larger than the leak conductance; thus it could significantly influence neuronal activity. This change in conductance may be due to stimulation of mechanoreceptors. Alternatively, US may perturb the lateral motion of phospholipids and produce nanopores, which then increase g us . These results may be important for understanding mechanisms underlying US-mediated modulation of neuronal activity and brain function. NEW & NOTEWORTHY We report a specific increase in membrane conductance produced by ultrasound (US) on neuronal membrane. When a 5-ms US tone burst was focused on a crayfish motor axon, it stochastically triggered either depolarization or a spike train. The depolarization was up to 60 mV in amplitude and 200 s in duration and therefore could significantly influence neuronal activity. Depolarization was still evoked by US burst in the presence of Na + and Ca 2+ channel blockers and had a reversal potential of -8.4 ± 2.3 mV, suggesting a nonselective permeability. US can be applied noninvasively in the form of a focused beam to deep brain areas through the skull and has been shown to modulate brain activity. Understanding the depolarization reported here should be helpful for improving the use of US for noninvasive modulation and stimulation in brain-related disease.

Abstract via europepmc.

SpeciesProcambarus clarkii (crayfish)
Subjects6, 11, 5swept preparations
Sessions per subjectnot applicable
Randomisednot applicable
Blindingnot applicable
Sham / controlnone
Auditory controlnot reported
Readout timingonline
Anaesthesianot applicable
Readoutsinvasive electrophysiologyTwo-electrode intracellular current-clamp recording of membrane potential and input resistance in crayfish motor axon; extracellular loose-patch recording in a subset
Direction of effectbidirectionalA focused ultrasound burst consistently produced a fast hyperpolarization, which was stochastically followed or superseded by subthreshold depolarization or action potentials; depolarization persisted after blocking voltage-gated Na+, K+ and HCN channels and was associated with decreased membrane resistance.
Adverse eventsnot applicable

Exposures

Exposure 1: Focused ultrasound burst applied to isolated crayfish motor axon

Target: crayfish neurons, neuromuscular junction — “motor axon of the crayfish opener neuromuscular junction
Device: custom-built

Pulse timing
Waveformcontinuous
Fundamental frequency (kHz)2,100✓✓
Pulse duration (ms)not applicable
Pulse repetition frequency (Hz)not applicable
Duty cycle (%)not applicable
Sonication duration (s)0.005✓✓
Pressure and intensity, by domain
Free-field pressure (kPa)100, 1,000swept✓✓
Free-field Isppa (W/cm²)not reported
Free-field Ispta (W/cm²)0.0042, 0.0094swept✓✓
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

Unless otherwise specified, responses were evoked by a single 5-ms, 2.1-MHz US tone burst focused (1-mm focal diameter) on the axon. US stimuli were delivered in 20-s cycles, each cycle consisting of a 10-s recorded trace followed by a 10-s interval before the next recording, over 2-4 h recording sessions; some protocols delivered bursts repeatedly at a low rate (about every 10 s) within a session.

Flags from extraction

  • n_subjectsQuote covers Groups 2 and 3 (n=11, n=5); Group 1 (n=6) is described in the same paragraph with garbled source formatting and is included in the list but not separately quoted.
  • exposures[0].free_field.pressure_kpa0.1-1 MPa is the overall pressure range used across experiments (from the abstract); individual figures cite specific intensities (e.g. 2.7-9.4 mW/cm2) for particular trials that are not individually itemised here.