Corticogeniculate neurons, corticotectal neurons, and suspected interneurons in visual cortex of awake rabbits: receptive-field properties, axonal properties, and effects of EEG arousal

1987 ◽  
Vol 57 (4) ◽  
pp. 977-1001 ◽  
Author(s):  
H. A. Swadlow ◽  
T. G. Weyand
Keyword(s):  
Electrical Stimulation ◽  
Visual Cortex ◽  
Receptive Field ◽  
Receptive Fields ◽  
Spontaneous Firing ◽  
Visual Responses ◽  
Firing Rates ◽  
Axonal Conduction ◽  
Conduction Velocities ◽  
Stimulation Of

The intrinsic stability of the rabbit eye was exploited to enable receptive-field analysis of antidromically identified corticotectal (CT) neurons (n = 101) and corticogeniculate (CG) neurons (n = 124) in visual area I of awake rabbits. Eye position was monitored to within 1/5 degrees. We also studied the receptive-field properties of neurons synaptically activated via electrical stimulation of the dorsal lateral geniculate nucleus (LGNd). Whereas most CT neurons had either complex (59%) or motion/uniform (15%) receptive fields, we also found CT neurons with simple (9%) and concentric (4%) receptive fields. Most complex CT cells were broadly tuned to both stimulus orientation and velocity, but only 41% of these cells were directionally selective. We could elicit no visual responses from 6% of CT cells, and these cells had significantly lower conduction velocities than visually responsive CT cells. The median spontaneous firing rates for all classes of CT neurons were 4-8 spikes/s. CG neurons had primarily simple (60%) and concentric (9%) receptive fields, and none of these cells had complex receptive fields. CG simple cells were more narrowly tuned to both stimulus orientation and velocity than were complex CT cells, and most (85%) were directionally selective. Axonal conduction velocities of CG neurons (mean = 1.2 m/s) were much lower than those of CT neurons (mean = 6.4 m/s), and CG neurons that were visually unresponsive (23%) had lower axonal conduction velocities than did visually responsive CG neurons. Some visually unresponsive CG neurons (14%) responded with saccadic eye movements. The median spontaneous firing rates for all classes of CG neurons were less than 1 spike/s. All neurons synaptically activated via LGNd stimulation at latencies of less than 2.0 ms had receptive fields that were not orientation selective (89% motion/uniform, 11% concentric), whereas most cells with orientation-selective receptive fields had considerably longer synaptic latencies. Most short-latency motion/uniform neurons responded to electrical stimulation of the LGNd (and visual area II) with a high-frequency burst (500-900 Hz) of three or more spikes. Action potentials of these neurons were of short duration, thresholds of synaptic activation were low, and spontaneous firing rates were the highest seen in rabbit visual cortex. These properties are similar to those reported for interneurons in several regions in mammalian central nervous system. Nonvisual sensory stimuli that resulted in electroencephalographic arousal (hippocampal theta activity) had a profound effect on the visual responses of many visual cortical neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Efferent neurons and suspected interneurons in motor cortex of the awake rabbit: axonal properties, sensory receptive fields, and subthreshold synaptic inputs

1994 ◽  
Vol 71 (2) ◽  
pp. 437-453 ◽  
Author(s):  
H. A. Swadlow
Keyword(s):  
Motor Cortex ◽  
Receptive Field ◽  
Receptive Fields ◽  
Sensory Stimulation ◽  
Spontaneous Firing ◽  
Firing Rates ◽  
Axonal Conduction ◽  
Conduction Velocities ◽  
Efferent Neurons ◽  
Stimulation Of

1. Properties of antidromically identified efferent neurons within the cortical representation of the vibrissae, sinus hairs, and philtrum were examined in motor cortex of fully awake adult rabbits. Efferent neurons were tested for both receptive field and axonal properties and included callosal (CC) neurons (n = 31), ipsilateral corticocortical (C-IC) neurons (n = 34) that project to primary somatosensory cortex (S-1), and corticofugal neurons of layer 5 (CF-5) (n = 33) and layer 6 (CF-6) (n = 32) that project to and/or beyond the thalamus. Appropriate collision tests demonstrated that substantial numbers of corticocortical efferent neurons project an axon to both the corpus callosum and to ipsilateral S-1. 2. Suspected interneurons (SINs, n = 37) were also studied. These neurons were not activated antidromically from any stimulus site but did respond synaptically to electrical stimulation of the ventrolateral (VL) thalamus and/or S-1 with a burst of three or more spikes at frequencies from 600 to > 900 Hz. All of these neurons also responded synaptically to stimulation of the corpus callosum. The action potentials of these neurons were much shorter in duration (mean = 0.48 ms), than those of efferent neurons (mean = 0.90 ms). 3. CF-5 neurons differed from CC, C-IC, and CF-6 neurons in their spontaneous firing rates, axonal properties, and receptive field properties. Whereas CF-5 neurons had a mean spontaneous firing rate of 4.1 spikes/s, CC, C-IC, and CF-6 neurons all had mean values of < 1 spike/s. Axonal conduction velocities of CF-5 neurons were much higher (mean = 12.76 m/s) than either CC (1.47 m/s), C-IC (0.97 m/s), or CF-6 (mean = 1.96 m/s) neurons. A decrease in antidromic latency (the "supernormal" period) followed a single prior impulse in most CC, C-IC, and CF-6 neurons but was minimal or absent in CF-5 neurons. Although all but two CF-5 neurons responded to peripheral sensory stimulation, many CC (35%), C-IC (59%), or CF-6 (66%) neurons did not. CC, CF-5, and CF-6 neurons that did not respond to sensory stimulation had significantly lower axonal conduction velocities and spontaneous firing rates than those that responded to such stimulation. 4. Sensory receptive fields of neurons in motor cortex were considerably larger than those observed in S-1 but were similar in size to those seen in secondary somatosensory cortex (S-2).(ABSTRACT TRUNCATED AT 400 WORDS)

Efferent neurons and suspected interneurons in binocular visual cortex of the awake rabbit: receptive fields and binocular properties

1988 ◽  
Vol 59 (4) ◽  
pp. 1162-1187 ◽  
Author(s):  
H. A. Swadlow
Keyword(s):  
Receptive Field ◽  
Receptive Fields ◽  
Visual Area ◽  
Orientation Tuning ◽  
Spontaneous Firing ◽  
Firing Rates ◽  
Axonal Conduction ◽  
Conduction Velocities ◽  
Efferent Neurons ◽  
Stimulation Of

1. In fully awake rabbits the stability of the two eyes was monitored and was sufficient to enable receptive-field analysis of antidromically identified efferent neurons and suspected interneurons in the binocular segment of visual area 1. Efferent neurons analyzed included callosal efferent neurons (CC neurons, n = 52), neurons projecting to visual area 2 (CV2 neurons, n = 35), corticotectal neurons (CT neurons, n = 43), and corticogeniculate neurons (CG neurons, n = 51). Six additional neurons projected a branching axon to both the corpus callosum and visual area 2. 2. Most CC and CV2 neurons were found in layer 2-3 and had receptive fields of the simple type. Only two corticocortical neurons with complex receptive fields were found. Orientation tuning ranges of CC and CV2 simple cells were similar and end stopping was prevalent in both CC (62%) and CV2 (45%) neurons. Axonal conduction velocities of CC and CV2 neurons were low (mean = 3.5 and 1.4 m/s, respectively) and visually nonresponsive CC neurons (19%) had conduction velocities that were significantly lower than visually responsive neurons. Spontaneous firing rates of corticocortical neurons were low (mean less than 1 spike/s) and these neurons responded to a lower range of stimulus velocities than did corticofugal neurons. 3. Most CG neurons had simple receptive fields and none had a complex field. Orientation tuning ranges of these neurons were comparable to those of CC and CV2 neurons, but a significantly smaller proportion (12%) were end stopped. Both spontaneous firing rates (mean = less than 1 spike/s) and axonal conduction velocities (mean = 2.4 m/s) of CG neurons were low and, as was found for CC neurons, visually nonresponsive CG neurons (25%) had significantly lower conduction velocities than did visually responsive neurons. 4. CT neurons had receptive fields that were predominantly complex (37%), motion/uniform (28%), or simple (26%). Conduction velocities (mean = 10.9 m/s) and spontaneous firing rates (mean = 7 spikes/s) of CT neurons of all receptive-field types were much higher than those of CC, CV2, and CG neurons. 5. An additional class of neurons was studied that responded synaptically at a short latency to electrical stimulation of the dorsal lateral geniculate nucleus (LGNd) with a burst of three or more spikes at frequencies of 600-900 Hz. These neurons showed a high degree of synaptic convergence, also responding synaptically with a high-frequency burst of spikes to stimulation of both visual area 2 and the corpus callosum.(ABSTRACT TRUNCATED AT 400 WORDS)

Efferent neurons and suspected interneurons in S-1 forelimb representation of the awake rabbit: receptive fields and axonal properties

1990 ◽  
Vol 63 (6) ◽  
pp. 1477-1498 ◽  
Author(s):  
H. A. Swadlow
Keyword(s):  
Receptive Field ◽  
Receptive Fields ◽  
Sensory Stimulation ◽  
Spontaneous Firing ◽  
Firing Rates ◽  
Axonal Conduction ◽  
Receptive Field Properties ◽  
Conduction Velocities ◽  
Efferent Neurons ◽  
Stimulation Of

1. Receptive-field properties of antidromically identified efferent neurons within the cutaneous forelimb representation of primary somatosensory cortex (S-1) were examined in fully awake rabbits. Efferent neurons studied included callosal neurons (CC neurons, n = 52), ipsilateral corticocortical neurons (C-IC neurons, n = 48) that project to or beyond the second somatosensory cortical area (S-2), and corticofugal neurons of layer 5 (CF-5 neurons, n = 97) and layer 6 (CF-6 neurons, n = 59) that project to and/or beyond the thalamus. 2. An additional class of neurons was studied that was not activated antidromically from any stimulus site, but which responded synaptically to electrical stimulation of the ventrobasal (VB) thalamus with a burst of three or more spikes at frequencies of 600 to greater than 900 Hz. Most of these neurons also responded synaptically to stimulation of S-2 and the corpus callosum. The action potentials of these neurons were much shorter (mean = 0.45 ms) than those of efferent neurons (mean = 0.95 ms). Such properties have been associated with interneurons found throughout the central nervous system, and these neurons are thereby referred to as suspected interneurons (SINs). 3. CF-5 neurons differed from CC, C-IC, and CF-6 neurons in their spontaneous firing rates, axonal properties, and receptive-field properties. Whereas CF-5 neurons had a mean spontaneous firing rate of 5.5 spikes/s, CC, C-IC, and CF-6 neurons had mean values of less than 1/s. Axonal conduction velocities of CF-5 neurons were much higher (mean = 12.92 m/s) than either CC (mean = 2.15 m/s), C-IC (mean = 1.31 m/s), or CF-6 (mean = 2.53 m/s) neurons. A decrease in antidromic latency (the "supernormal" period) that was dependent on prior impulse activity was seen in the great majority of CC, C-IC, and CF-6 neurons but was either minimal or absent in CF-5 neurons of comparable conduction velocity. A higher proportion of CF-5 neurons (98%) responded to peripheral sensory stimulation than did either CC (75%), C-IC (71%), or CF-6 (51%) neurons. CF-6 and C-IC neurons that did not respond to sensory stimulation had significantly lower axonal conduction velocities and spontaneous firing rates than those that responded to such stimulation. 4. Cutaneous receptive fields were seen in most neurons that could be driven by peripheral stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)

Efferent neurons and suspected interneurons in second somatosensory cortex of the awake rabbit: receptive fields and axonal properties

1991 ◽  
Vol 66 (4) ◽  
pp. 1392-1409 ◽  
Author(s):  
H. A. Swadlow
Keyword(s):  
Receptive Field ◽  
Corpus Callosum ◽  
Somatosensory Cortex ◽  
Sensory Stimulation ◽  
Spontaneous Firing ◽  
Firing Rates ◽  
Axonal Conduction ◽  
Conduction Velocities ◽  
Efferent Neurons ◽  
Stimulation Of

1. Receptive-field properties of antidromically identified efferent neurons within the representation of vibrissae and sinus hairs above the mouth were examined in secondary somatosensory cortex (S-2) of fully awake adult rabbits. Efferent neurons studied included callosal neurons (CC neurons, n = 88), ipsilateral corticocortical neurons (C-IC neurons, n = 51) that project to primary somatosensory cortex (S-1), and corticofugal neurons of layer 5 (CF-5 neurons, n = 63) and layer 6 (CF-6 neurons, n = 42) that project to and/or beyond the thalamus. Appropriate collision tests demonstrated that substantial numbers of corticocortical efferent neurons (21 of 113 tested) project an axon to both the corpus callosum and to ipsilateral S-1. 2. Suspected interneurons (SINs, n = 62) were also studied. These neurons were not activated antidromically from any stimulus site but did respond synaptically to electrical stimulation of the ventrobasal (VB) thalamus with a burst of three or more spikes at frequencies of 600 to greater than 900 Hz. Most of these neurons also responded synaptically to stimulation of S-1 and the corpus callosum. The action potentials of these neurons were much shorter (mean, 0.49 ms) than those of efferent neurons (mean, 1.01 ms). 3. CF-5 neurons differed from CC, C-IC, and CF-6 neurons in their spontaneous firing rates, axonal properties, and receptive-field properties. Whereas CF-5 neurons had a mean spontaneous firing rate of 5.7 spikes/s, CC, C-IC, and CF-6 neurons all had mean values of less than 1/s. Axonal conduction velocities of CF-5 neurons were much higher (mean, 11.90 m/s) than either CC (mean, 2.63 m/s), C-IC (mean, 0.86 m/s), or CF-6 (mean, 1.73 m/s) neurons. A decrease in antidromic latency (the "supernormal" period), which was dependent on prior impulse activity, was seen in most CC, C-IC, and CF-6 neurons but was minimal or absent in CF-5 neurons of comparable conduction velocity. Although all CF-5 neurons responded to peripheral sensory stimulation, many CC (52%), C-IC (49%), and CF-6 (55%) neurons did not. CC and CF-6 neurons that did not respond to sensory stimulation had significantly lower axonal conduction velocities and spontaneous firing rates than those that responded to such stimulation. Whereas no CC, C-IC, or CF-6 neuron responded synaptically to callosal stimulation, 43% of CF-5 neurons (and 78% of SINs) did so respond. Similar differences in synaptic responsivity to stimulation of S-1 were seen in these populations.(ABSTRACT TRUNCATED AT 400 WORDS)

scholarly journals Neural Mechanisms of Spatial Selective Attention in Areas V1, V2, and V4 of Macaque Visual Cortex

1997 ◽  
Vol 77 (1) ◽  
pp. 24-42 ◽  
Author(s):  
Steven J. Luck ◽  
Leonardo Chelazzi ◽  
Steven A. Hillyard ◽  
Robert Desimone
Keyword(s):  
Visual Cortex ◽  
Receptive Field ◽  
Selective Attention ◽  
Receptive Fields ◽  
Simultaneous Presentation ◽  
Neural Mechanisms ◽  
Spontaneous Firing ◽  
Firing Rates ◽  
Behavioral Paradigm ◽  
Sensory Responses

Luck, Steven J., Leonardo Chelazzi, Steven A. Hillyard, and Robert Desimone. Neural mechanisms of spatial selective attention in areas V1, V2, and V4 of macaque visual cortex. J. Neurophysiol. 77: 24–42, 1997. Many neurons in extrastriate visual cortex have large receptive fields, and this may lead to significant computational problems whenever multiple stimuli fall within a single field. Previous studies have suggested that when multiple stimuli fall within a cell's receptive field, they compete for the cell's response in a manner that can be biased in favor of attended stimuli. In the present study we examined this role of attention in areas V1, V2, and V4 of macaque monkeys with the use of a behavioral paradigm in which attention was directed to one of two stimulus locations. When two stimuli were presented simultaneously inside the cell's receptive field (which could be accomplished only in areas V2 and V4), we found that the cell's response was strongly influenced by which of the two stimuli was attended. The size of this attention effect was reduced when the attended and ignored stimuli were presented sequentially rather than simultaneously. In addition, the effects became very weak and inconsistent in these areas when only one of the two stimuli was located inside the receptive field. Attention thus modulated sensory responses primarily when two or more simultaneous stimuli competed for access to a neuron's receptive field. As in areas V2 and V4, attention did not modulate sensory responses in area V1 when only a single stimulus was inside the receptive field. In addition, the small receptive fields in this area precluded the simultaneous presentation of attended and ignored stimuli inside the receptive field, making it impossible to determine whether attention effects would be observed under the conditions that led to consistent attention effects in areas V2 and V4. Spontaneous firing rates in areas V2 and V4 were found to be 30–40% higher when attention was directed inside rather than outside the receptive field, even when no stimulus was present in the receptive field. Spontaneous firing rates also varied according to the particular location within the receptive field that was attended. These shifts in spontaneous activity may reflect a top-down signal that biases responses in favor of stimuli at the attended location.

Effects of norepinephrine on the activity of visual neurons in the superior colliculus of the hamster

1999 ◽  
Vol 16 (3) ◽  
pp. 541-555 ◽  
Author(s):  
YI ZHANG ◽  
RICHARD D. MOONEY ◽  
ROBERT W. RHOADES
Keyword(s):  
Electrical Stimulation ◽  
Visual Cortex ◽  
Superior Colliculus ◽  
Visual Stimulation ◽  
Optic Chiasm ◽  
Evoked Responses ◽  
Visual Responses ◽  
Signal To Noise ◽  
Direct Stimulation ◽  
Stimulation Of

Single-unit recording and micropressure ejection techniques were used to test the effects of norepinephrine (NE) on the responses of neurons in the superficial layers (the stratum griseum superficiale and stratum opticum) of the hamster's superior colliculus (SC). Application of NE suppressed visually evoked responses by ≥30% in 75% of 40 neurons tested and produced ≥30% augmentation of responses in only 5%. The decrement in response strength was mimicked by application of the α2 adrenoceptor agonist, p-aminoclonidine, the nonspecific β agonist, isoproterenol, and the β1 agonist, dobutamine. These agents had similar effects on responses evoked by electrical stimulation of the optic chiasm and visual cortex. The α1 agonist, methoxamine, augmented the light-evoked responses of 53% of 49 SC cells by ≥30%, but had little effect on responses evoked by electrical stimulation of optic chiasm or visual cortex. The effects of adrenergic agonists upon the glutamate-evoked responses of SC cells that were synaptically “isolated” by concurrent application of Mg2+ were similar to those obtained during visual stimulation. Analysis of effects of NE on visually evoked and background activity indicated that application of this amine did not significantly enhance signal-to-noise ratios for most superficial layer SC neurons, and signal-to-noise ratios were in some cases reduced. These results indicate that NE acts primarily through α2 and β1 receptors to suppress the visual responses of SC neurons. Activation of either of these receptors reduces the responses of SC neurons to either of their two major visual inputs as well as to direct stimulation by glutamate, and it would thus appear that these effects are primarily postsynaptic.

Binocular convergence of excitatory and inhibitory synaptic pathways onto neurons of cat visual cortex

1990 ◽  
Vol 4 (6) ◽  
pp. 625-629 ◽  
Author(s):  
David Ferster
Keyword(s):  
Electrical Stimulation ◽  
Visual Cortex ◽  
Receptive Field ◽  
Synaptic Input ◽  
Intracellular Recording ◽  
Cortical Neurons ◽  
Field Studies ◽  
The Other ◽  
Synaptic Potential ◽  
Stimulation Of

AbstractWhen a cortical neuron receives synaptic input from both eyes, do the synaptic pathways that mediate the input from each eye match? In this study, inputs from the two eyes were compared by measuring the latencies of EPSPs and IPSPs evoked by electrical stimulation of the two optic nerves. For binocular neurons, these latencies invariably matched closely, indicating that the pathways from the two eyes contain the same number of synapses; monosynaptic input from lamina A of the lateral geniculate nucleus (LGN) is always matched by monosynaptic input from lamina A1. Conversely, polysynaptic input from one eye, either excitatory or inhibitory, is invariably accompanied by similar input from the other eye. In addition, the match between the two eyes in latency indicates that for each eye a synaptic potential is mediated by the same type of afferent, either X or Y.Judging from intracellular recording, 75% of the neurons studied were binocular, that is, EPSPs could be evoked from either eye. In the remaining 25%, EPSPs could be evoked from only one eye, in agreement with extracellular receptive field studies in which 30% of cortical neurons are monocular.

Mechanoafferent neurons innervating tail of Aplysia. I. Response properties and synaptic connections

1983 ◽  
Vol 50 (6) ◽  
pp. 1522-1542 ◽  
Author(s):  
E. T. Walters ◽  
J. H. Byrne ◽  
T. J. Carew ◽  
E. R. Kandel
Keyword(s):  
Electrical Stimulation ◽  
Receptive Field ◽  
Motor Neurons ◽  
Action Potentials ◽  
Receptive Fields ◽  
The Body ◽  
Tail Region ◽  
Withdrawal Reflex ◽  
Somatotopic Organization ◽  
Stimulation Of

Mechanical, chemical, or electrical stimulation of the tail elicits a short-latency (less than 1 s) tail-withdrawal reflex that is graded with the intensity of the stimulus. The tail-withdrawal reflex is not elicited by stimulation of parts of the body outside of the tail region. Mechanoafferent neurons innervating the tail are located in a small subcluster within a large, homogeneous group of medium-size (40-80 micron) cells on the ventrocaudal (VC) surface of each pleural ganglion. The tail sensory neurons within this large VC cluster are activated by tactile pressure or by electrical stimulation of discrete regions of the tail. They adapt slowly to maintained stimulation and sometimes respond to stimulus offset as well. Both mechanical and electrical stimuli produce responses that are graded with the intensity of the stimulus. Cells in the VC cluster appear to be primary mechanoreceptors because they have axons in peripheral nerves (including nerves innervating the tail), they exhibit action potentials lacking prepotentials in response to tactile stimulation, and these action potentials are still produced by cutaneous stimulation when peripheral and central chemical synaptic transmission is blocked. Stimulation of fields all over the body surface evokes synaptically mediated hyperpolarizing responses in individual mechanoafferent neurons that may represent afferent inhibition. Hyperpolarizing responses lasting many seconds can be produced by brief cutaneous stimuli. The mechanoafferent neurons innervating the tail region make strong monosynaptic connections to tail motor neurons in the ipsilateral pedal ganglion, and through these connections this subpopulation of the VC neurons appears to make a substantial contribution to the short-latency tail-withdrawal reflex. In addition, the combined excitatory receptive fields of these mechanoafferents match the excitatory receptive field of the tail-withdrawal reflex. Mechanoafferent neurons in the VC cluster that have receptive fields on other parts of the body (outside the excitatory receptive field of the tail-withdrawal reflex) have not been observed to make monosynaptic connections to the tail motor neurons. The neurons innervating the tail are reliably found in a discrete region within the larger VC cluster. In addition to this gross somatotopic organization, there is evidence of a finer level of somatotopic organization between the position of the excitatory receptive field on the tail and the position of the cell soma in the tail subcluster.(ABSTRACT TRUNCATED AT 400 WORDS)

scholarly journals Electrical stimulation of the nucleus basalis of meynert: a systematic review of preclinical and clinical data

2021 ◽  
Vol 11 (1) ◽  
Author(s):  
Muhammad Nazmuddin ◽  
Ingrid H. C. H. M. Philippens ◽  
Teus van Laar
Keyword(s):  
Electrical Stimulation ◽  
Clinical Data ◽  
Animal Studies ◽  
Receptive Fields ◽  
Lewy Body Dementia ◽  
Clinical Situation ◽  
Nucleus Basalis ◽  
Nucleus Basalis Of Meynert ◽  
Clinical Effects ◽  
Stimulation Of

AbstractDeep brain stimulation (DBS) of the nucleus basalis of Meynert (NBM) has been clinically investigated in Alzheimer’s disease (AD) and Lewy body dementia (LBD). However, the clinical effects are highly variable, which questions the suggested basic principles underlying these clinical trials. Therefore, preclinical and clinical data on the design of NBM stimulation experiments and its effects on behavioral and neurophysiological aspects are systematically reviewed here. Animal studies have shown that electrical stimulation of the NBM enhanced cognition, increased the release of acetylcholine, enhanced cerebral blood flow, released several neuroprotective factors, and facilitates plasticity of cortical and subcortical receptive fields. However, the translation of these outcomes to current clinical practice is hampered by the fact that mainly animals with an intact NBM were used, whereas most animals were stimulated unilaterally, with different stimulation paradigms for only restricted timeframes. Future animal research has to refine the NBM stimulation methods, using partially lesioned NBM nuclei, to better resemble the clinical situation in AD, and LBD. More preclinical data on the effect of stimulation of lesioned NBM should be present, before DBS of the NBM in human is explored further.

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