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Low-intensity ultrasound directly modulates neural activity of the cerebellar cortex

Ruo-Shui Xu, Xue-Mei Wu, Zhi-Qi Xiong

Brain Stimulation 2023, 16, 918-926 · 10.1016/j.brs.2023.05.012

rodenthealthyothercellular imagingbehaviourhistology molecular

Abstract

Background Low-intensity ultrasound is a noninvasive neuromodulation technique with the potential to focally manipulate deep brain activity at millimeter-scale resolution. However, there have been controversies over the direct influence of ultrasound on neurons, due to an indirect auditory activation. Besides, the capacity of ultrasound to stimulate the cerebellum remains underestimated. Objective To validate the direct neuromodulation effects of ultrasound on the cerebellar cortex from both cellular and behavioral levels. Methods Two-photon calcium imaging were used to measure the neuronal responses of cerebellar granule cells (GrCs) and Purkinje cells (PCs) to ultrasound application in awake mice. And a mouse model of paroxysmal kinesigenic dyskinesia (PKD), in which direct activation of the cerebellar cortex leads to dyskinetic movements, was used to assess the ultrasound-induced behavioral responses. Results Low-intensity ultrasound stimulus (0.1 W/cm 2 ) evoked rapidly increased and sustained neural activity in GrCs and PCs at targeted region, while no significant changes in calcium signals were observed responding to off-target stimulus. The efficacy of ultrasonic neuromodulation relies on acoustic dose modified by ultrasonic duration and intensity. In addition, transcranial ultrasound reliably triggered dyskinesia attacks in proline-rich transmembrane protein 2 (Prrt2) mutant mice, suggesting that the intact cerebellar cortex were activated by ultrasound. Conclusion Low-intensity ultrasound directly activates the cerebellar cortex in a dose-dependent manner, and thus serves as a promising tool for cerebellar manipulation.

Abstract via europepmc.

Speciesmouse (C57BL/6J; Grin2c-iCre/Ai93D; Prrt2-mutant)
Subjectsnot reported animals
Sessions per subjectnot reported
Randomisednot reported
Blindingnot reported
Sham / controlactive control site
Auditory controlnot reported
Readout timingboth
Anaesthesiaawake
Readoutscellular imaging, behaviour, histology molecularTwo-photon calcium imaging of GCaMP6f in cerebellar granule cell boutons and Purkinje cell dendrites; dyskinesia behavioural scoring; TUNEL staining; anti-GFP immunohistochemistry
Direction of effectexcitatoryUltrasound rapidly increased and sustained calcium activity in cerebellar granule cell boutons and Purkinje cell dendrites at the targeted region in a dose (duration/intensity)-dependent manner, with no significant change to an off-target site; transcranial ultrasound to the cerebellar hemisphere also reliably triggered dyskinesia attacks in Prrt2-mutant mice.
Adverse eventsnone observedTUNEL staining after ultrasound stimulation did not reveal TUNEL-positive apoptotic signals, and no cellular damage was observed in animals in which ultrasound triggered dyskinesia/cerebellar spreading depolarization.

Exposures

Exposure 1: Cerebellar cortex GrC/PC imaging and dyskinesia induction (88 kHz, CW, 5-10 s)

Target: cerebellar cortex — “cerebellar cortex, lobule VI-VIII (granule cell boutons and Purkinje cell dendrites); cerebellar hemisphere (dyskinesia induction)
Device: other named manufacturer · Hainertec · HNN–4SS-0890

Pulse timing
Waveformcontinuous
Fundamental frequency (kHz)88✓✓
Pulse duration (ms)not applicable
Pulse repetition frequency (Hz)not applicable
Duty cycle (%)100?
Sonication duration (s)5, 10swept✓✓
Pressure and intensity, by domain
Free-field pressure (kPa)not reported
Free-field Isppa (W/cm²)0.1✓✓
Free-field Ispta (W/cm²)not reported
In-situ estimatenot reported
In-situ pressure (kPa)not reported
In-situ Isppa (W/cm²)not reported
In-situ Ispta (W/cm²)not reported
Protocol, in the paper’s words

Continuous-wave (CW) ultrasonic stimuli of 5 or 10 s at 0.1 W/cm^2 were delivered to the cerebellar cortex (imaging window over lobule VI-VIII) during two-photon calcium imaging of GrC boutons; for Purkinje cell dendrite imaging, 5-s pulses were delivered every 30 s for a total of 10 trials. For dyskinesia induction, transcranial CW ultrasound (10 s, 0.1 W/cm^2) was applied to the cerebellar hemisphere; an off-target stimulus (10 s, 0.1 W/cm^2, same pattern) was applied to dental cement over the cerebrum. Each stimulation pattern was repeated 5 times, interleaved with at least 8 h for recovery.

Exposure 2: Cerebellar cortex acoustic dose-response (127 kHz tone bursts, 80/500 ms)

Target: cerebellar cortex — “cerebellar cortex (Purkinje cell dendrites); cerebellar hemisphere (500-ms on-target dyskinesia test)
Device: custom-built

Pulse timing
Waveformcontinuous
Fundamental frequency (kHz)127✓✓
Pulse duration (ms)not applicable
Pulse repetition frequency (Hz)not applicable
Duty cycle (%)100?
Sonication duration (s)0.08, 0.5swept✓✓
Pressure and intensity, by domain
Free-field pressure (kPa)not reported
Free-field Isppa (W/cm²)0.14, 1.32, 4.4swept✓✓
Free-field Ispta (W/cm²)not reported
In-situ estimatenot reported
In-situ pressure (kPa)not reported
In-situ Isppa (W/cm²)not reported
In-situ Ispta (W/cm²)not reported
Protocol, in the paper’s words

Single uninterrupted tone bursts of 80 ms or 500 ms at 0.14, 1.32 or 4.4 W/cm^2 were delivered once per 20-s trial, for a total of 14-20 trials, to assess dose-dependence of Purkinje cell dendrite responses. A 500-ms, 4.4 W/cm^2 on-target burst was also tested for its ability to trigger dyskinesia in Prrt2-mutant mice.

Flags from extraction

  • n_subjectsMany separate sub-experiments (imaging, dose-response, dyskinesia) each report different, non-overlapping animal/session/bouton counts; no single study-wide total of animals exposed to ultrasound is stated.
  • n_sessions_per_subjectNumber of sessions per subject varies by sub-experiment (single imaging session vs. 5 repeated dyskinesia trials with recovery); no single value applies to the whole study.
  • exposures[1].fundamental_frequency_khzPaper states the second transducer was driven 'around 127 kHz', an approximate rather than exact value.
  • exposures[0].unspecified_domain.isppa_w_cm2Domain (free-field vs. in-brain) of the reported intensity is not stated in the main text; a figure legend indicates intensities were measured with a hydrophone at 3 mm from the horn surface (suggesting a free-field-like calibration), but this is not stated as such in running text.
  • exposures[1].unspecified_domain.isppa_w_cm2Domain (free-field vs. in-brain) of the reported intensities is not stated in the main text.
  • exposures[0].timing.duty_cycle_pctDuty cycle of 100% is inferred from the paper's designation of the stimulus as continuous wave (CW); not independently stated as a percentage.
  • exposures[1].timing.duty_cycle_pctDuty cycle of 100% is inferred from the single uninterrupted tone-burst-per-trial description; not independently stated as a percentage.