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Induction of mandibular tremor using electrolytic lesion of the ventrolateral striatum or using subchronic haloperidol therapy in male rats: An electromyographic comparison
Inducción del temblor mandibular por lesión electrolítica del estriado ventrolateral y por el tratamiento subcrónico con haloperidol en rata macho: un contraste electromiográfico
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Calibración horizontal: 200<span class="elsevierStyleHsp" style=""></span>ms. Calibración vertical: 1<span class="elsevierStyleHsp" style=""></span>mV. B) Amplitud de la actividad basal (mV) durante el registro EMG del músculo temporal en un grupo de ratas control (C), con haloperidol (H), con lesión electrolítica en el EVL (L).</p>" ] ] ] "autores" => array:1 [ 0 => array:2 [ "autoresLista" => "G. Herrera-Meza, J. Manzo, M.E. Hernández, M. Miquel, L.I. García" "autores" => array:5 [ 0 => array:2 [ "nombre" => "G." "apellidos" => "Herrera-Meza" ] 1 => array:2 [ "nombre" => "J." "apellidos" => "Manzo" ] 2 => array:2 [ "nombre" => "M.E." "apellidos" => "Hernández" ] 3 => array:2 [ "nombre" => "M." "apellidos" => "Miquel" ] 4 => array:2 [ "nombre" => "L.I." 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"apellidos" => "García" "email" => array:2 [ 0 => "luisgarcia@uv.mx" 1 => "garcial00@yahoo.com.mx" ] "referencia" => array:2 [ 0 => array:2 [ "etiqueta" => "<span class="elsevierStyleSup">a</span>" "identificador" => "aff0005" ] 1 => array:2 [ "etiqueta" => "<span class="elsevierStyleSup">*</span>" "identificador" => "cor1" ] ] ] ] "afiliaciones" => array:3 [ 0 => array:3 [ "entidad" => "Centro de Investigaciones Cerebrales, Universidad Veracruzana, Xalapa, Veracruz, Mexico" "etiqueta" => "a" "identificador" => "aff0005" ] 1 => array:3 [ "entidad" => "Instituto de Neuroetología, Universidad Veracruzana, Xalapa, Veracruz, Mexico" "etiqueta" => "b" "identificador" => "aff0010" ] 2 => array:3 [ "entidad" => "Departamento de Psicobiología, Universidad de Jaume I, Castellón, Spain" "etiqueta" => "c" "identificador" => "aff0015" ] ] "correspondencia" => array:1 [ 0 => array:3 [ "identificador" => "cor1" "etiqueta" => "⁎" "correspondencia" => "Corresponding author." ] ] ] ] "titulosAlternativos" => array:1 [ "es" => array:1 [ "titulo" => "Inducción del temblor mandibular por lesión electrolítica del estriado ventrolateral y por el tratamiento subcrónico con haloperidol en rata macho: un contraste electromiográfico" ] ] "resumenGrafico" => array:2 [ "original" => 0 "multimedia" => array:7 [ "identificador" => "fig0010" "etiqueta" => "Figure 2" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr2.jpeg" "Alto" => 975 "Ancho" => 1360 "Tamanyo" => 29564 ] ] "descripcion" => array:1 [ "en" => "<p id="spar0050" class="elsevierStyleSimplePara elsevierViewall">Mean value<span class="elsevierStyleHsp" style=""></span>±<span class="elsevierStyleHsp" style=""></span>SD of the amplitude (mV) of EMG activity in the temporal muscle at baseline in the control group, haloperidol group, and electrolytic VLS lesion group.</p>" ] ] ] "textoCompleto" => "<span class="elsevierStyleSections"><span id="sec0005" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0065">Introduction</span><p id="par0005" class="elsevierStylePara elsevierViewall">Onset of mandibular tremor or tremulous jaw movement (TJM) in rats has been linked to changes in striatal mechanisms.<a class="elsevierStyleCrossRefs" href="#bib0005"><span class="elsevierStyleSup">1,2</span></a> Most TJM cases are induced by changes in striatal dopaminergic neurotransmission, which in turn are caused by dopaminergic antagonism<a class="elsevierStyleCrossRefs" href="#bib0015"><span class="elsevierStyleSup">3–5</span></a> or by the effect of dopamine depletion in the ventrolateral striatum elicited by 6-hydroxydopamine (6-OHDA). Both conditions lead to TJMs.</p><p id="par0010" class="elsevierStylePara elsevierViewall">The lateral part of the striatum is associated with motor and sensorimotor function; the ventrolateral area in particular is involved in motor control of the mouth area and limbs.<a class="elsevierStyleCrossRefs" href="#bib0040"><span class="elsevierStyleSup">8,9</span></a> As a result, the ventrolateral part of the striatum is specifically implicated in TJM.<a class="elsevierStyleCrossRef" href="#bib0035"><span class="elsevierStyleSup">7</span></a> That area receives dopaminergic pathways from the compact substantia nigra,<a class="elsevierStyleCrossRefs" href="#bib0050"><span class="elsevierStyleSup">10,11</span></a> which contains both neurotransmitters and neuromodulators such as ACh and DA.<a class="elsevierStyleCrossRef" href="#bib0060"><span class="elsevierStyleSup">12</span></a></p><p id="par0015" class="elsevierStylePara elsevierViewall">TJM characteristically occurs in phasic bursts of repetitive jaw movements that are not triggered by any particular stimulus<a class="elsevierStyleCrossRef" href="#bib0065"><span class="elsevierStyleSup">13</span></a> and have a frequency distribution of 3 to 7<span class="elsevierStyleHsp" style=""></span>Hz.<a class="elsevierStyleCrossRef" href="#bib0070"><span class="elsevierStyleSup">14</span></a> As a result, the neurochemical, anatomical, and EMG findings in rodent TJM are comparable with tremor recorded in parkinsonian patients,<a class="elsevierStyleCrossRefs" href="#bib0065"><span class="elsevierStyleSup">13–15</span></a> and TJM associated with the manipulation of dopamine in striatal mechanisms therefore provides a functional rat model for analysing certain changes in basal ganglia.</p><p id="par0020" class="elsevierStylePara elsevierViewall">In this context, and given evidence showing that the ventrolateral striatum is the area directly related to the origin of TJM, we hypothesise that electrolytic lesion to the VLS will cause jaw movements that differ electromyographically from those derived from dopaminergic antagonism by means of subchronic (14-day) treatment with IP haloperidol. The purpose of this study was to describe electromyographic readings from TJMs generated by bilateral electrolytic lesion to the VLS and to compare them to TJMs induced by subchronic treatment with IP haloperidol in Wistar rats.</p></span><span id="sec0010" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0070">Materials and methods</span><span id="sec0015" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0075">Experimental subjects</span><p id="par0025" class="elsevierStylePara elsevierViewall">Twenty-two male rats ranging in weight from 250 to 350<span class="elsevierStyleHsp" style=""></span>g were housed individually in transparent acrylic cages measuring 44<span class="elsevierStyleHsp" style=""></span>cm<span class="elsevierStyleHsp" style=""></span>×<span class="elsevierStyleHsp" style=""></span>34<span class="elsevierStyleHsp" style=""></span>cm<span class="elsevierStyleHsp" style=""></span>×<span class="elsevierStyleHsp" style=""></span>20<span class="elsevierStyleHsp" style=""></span>cm, each of which contained 5<span class="elsevierStyleHsp" style=""></span>cm sterile sawdust bedding (Harlan, Mexico). Rats were kept with ad libitum feeding and a 12<span class="elsevierStyleHsp" style=""></span>×<span class="elsevierStyleHsp" style=""></span>12<span class="elsevierStyleHsp" style=""></span>hour inverted light–dark cycle (lights were turned on at 20.00). All experiments were carried out in strict accordance with official Mexican guidelines for the care and use of laboratory animals (NOM-062-ZOO-1999) and <span class="elsevierStyleItalic">Guide for the Care and Use of Laboratory Animals</span> (NIH, Washington D.C., USA).</p></span><span id="sec0020" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0080">Study groups</span><p id="par0030" class="elsevierStylePara elsevierViewall">Rats were assigned at random to 3 different groups. The first group (<span class="elsevierStyleItalic">n</span><span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>8) received IP haloperidol during 14 days (1.5<span class="elsevierStyleHsp" style=""></span>mg/kg) and the second group (<span class="elsevierStyleItalic">n</span><span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>8) received IP saline vehicle (1.5<span class="elsevierStyleHsp" style=""></span>ml/kg) during the same period. The haloperidol dose and treatment regimen were calculated based on the descriptions by Salamone et al.<a class="elsevierStyleCrossRef" href="#bib0080"><span class="elsevierStyleSup">16</span></a> and Trevitt et al.<a class="elsevierStyleCrossRef" href="#bib0085"><span class="elsevierStyleSup">17</span></a> The third group (<span class="elsevierStyleItalic">n</span><span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>6) consisted of animals subjected to a bilateral electrolytic lesion in the VLS.</p><p id="par0035" class="elsevierStylePara elsevierViewall">After 14 days, all rats treated with haloperidol (RBI Research Biochemicals International, Natick, MA, 01760, USA) exhibited TJM. The lesion group presented the same response at 24<span class="elsevierStyleHsp" style=""></span>hours; a 10-minute video recording of the response was made for each study group prior to EMG testing. Subsequently, only 2 rats from each group were selected at random to undergo electromyographic measurement of TJM and contrast responses to pharmacological and electrolytic changes. Rats with VLS lesion were treated with flunixin meglumine as an analgesic (2.5<span class="elsevierStyleHsp" style=""></span>mg/kg) and enrofloxacin as an antibiotic (5<span class="elsevierStyleHsp" style=""></span>mg/kg) in addition to subcutaneous fluid therapy with an electrolyte solution (Hartmann's solution 0.9%).</p></span><span id="sec0025" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0085">Electrolytic lesion to the ventrolateral striatum</span><p id="par0040" class="elsevierStylePara elsevierViewall">Rats were anaesthetised with mixture of ketamine (60<span class="elsevierStyleHsp" style=""></span>mg/kg, Bayer) and xylazine (8<span class="elsevierStyleHsp" style=""></span>mg/kg, Pisa). While animals were under deep anaesthesia, researchers drilled burr holes using the following stereotaxic coordinates<a class="elsevierStyleCrossRef" href="#bib0090"><span class="elsevierStyleSup">18</span></a>: anterior–posterior (AP)<span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>8.74<span class="elsevierStyleHsp" style=""></span>mm anterior to lambda, lateral (L)<span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>±4.4<span class="elsevierStyleHsp" style=""></span>mm, and ventral (V)<span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>3.40<span class="elsevierStyleHsp" style=""></span>mm from the interaural line. Using a stereotaxic instrument (Stoelting Co, USA), researchers then placed a single shaft stainless steel microelectrode 250<span class="elsevierStyleHsp" style=""></span>μm in diameter (Stoelting Co, USA). The electrode delivered 2.5<span class="elsevierStyleHsp" style=""></span>mA direct anodal current during 30<span class="elsevierStyleHsp" style=""></span>seconds using a Grass model S48 stimulator (Astro-Med Inc., USA) connected in series to a Grass model CCU1 constant current unit (Astro-Med Inc., USA).</p></span><span id="sec0030" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0090">Implanting electrodes for electromyography recordings</span><p id="par0045" class="elsevierStylePara elsevierViewall">Rats in all 3 groups were fitted with bipolar silver wire electrodes (Grass EW10AG) in the right temporal muscle 24<span class="elsevierStyleHsp" style=""></span>hours before electromyography. These electrodes measured 254<span class="elsevierStyleHsp" style=""></span>μm in diameter and were approximately 3<span class="elsevierStyleHsp" style=""></span>cm long with a 3<span class="elsevierStyleHsp" style=""></span>mm exposed segment at the end. Electrode leads were routed subcutaneously towards the rostral part of the skull and cemented with dental acrylic (MDC Dental).</p></span><span id="sec0035" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0095">Recording tremulous jaw movement</span><p id="par0050" class="elsevierStylePara elsevierViewall">Researchers recorded waking EMG activity. Animals with VLS lesion were assessed 48<span class="elsevierStyleHsp" style=""></span>hours after lesion induction. Assessment of animals treated with haloperidol or saline vehicle took place on day 14 of treatment.</p><p id="par0055" class="elsevierStylePara elsevierViewall">Implanted electrodes were connected to a Grass 15A54, 15LT amplifier (Astro-Med Inc., USA) and simultaneously to a Grass AM9 audio monitor (Astro-Med Inc., USA). The amplified analogue signal was relayed through an interface to a PVA-16 PolyVIEW Adaptor Unit to be digitised and then to a PC to calculate and analyse each animal's responses using the Grass PolyVIEW Data Acquisition and Analysis System 16 v 1.0.</p></span><span id="sec0040" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0100">Perfusion</span><p id="par0060" class="elsevierStylePara elsevierViewall">After the EMG recording, all animals were injected with high-dose sodium pentobarbital anaesthesia (0.6<span class="elsevierStyleHsp" style=""></span>ml/kg; Sedalpharma, Mexico). Transcardial perfusion was then performed to extract the brain, which was then cryoprotected in graded sucrose concentrations (10%, 20%, and 30%) diluted in a 0.1<span class="elsevierStyleHsp" style=""></span>M phosphate buffer. Researchers used a cryostat at −24<span class="elsevierStyleHsp" style=""></span>°C (LEICA CM1850) to make coronal slices 60<span class="elsevierStyleHsp" style=""></span>μm thick and verify lesion location in those animals with lesions.</p></span><span id="sec0045" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0105">Statistical analysis</span><p id="par0065" class="elsevierStylePara elsevierViewall">Researchers prepared graphs of the distribution of amplitude frequencies and EMG activity frequencies in the temporal muscle at baseline and during TJM in rats and recorded modal values. We also created general linear models with a design of main and nested (hierarchical) factors as indicated for analyses with pseudoreplications<a class="elsevierStyleCrossRef" href="#bib0095"><span class="elsevierStyleSup">19</span></a>:<elsevierMultimedia ident="eq0005"></elsevierMultimedia></p><p id="par0070" class="elsevierStylePara elsevierViewall">Here, <span class="elsevierStyleItalic">y</span> is the response variable (frequency and amplitude), <span class="elsevierStyleItalic">G</span> is the group, <span class="elsevierStyleItalic">R</span> is the rat, and <span class="elsevierStyleItalic">Ps</span> is pseudoreplication of EMG trace. In this analysis, the same model was adjusted by the amplitude of baseline activity in all groups. Response variables were fitted to Poisson-distributed data with a log-link function. All analyses were performed using JMP 6 statistical software (SAS Inc, Cary NC, USA, 2005).</p></span></span><span id="sec0050" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0110">Results</span><span id="sec0055" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0115">Electromyography recordings of baseline activity</span><p id="par0075" class="elsevierStylePara elsevierViewall">Baseline temporal muscle activity in the control, haloperidol, and electrolytic VLS groups showed different response patterns as evidenced in the 1-second EMG trace (<a class="elsevierStyleCrossRef" href="#fig0005">Fig. 1</a>A). Amplitude in control group animals ranged from 0.2 to 0.3<span class="elsevierStyleHsp" style=""></span>mV, while amplitude in the haloperidol group ranged from 0.3 to 0.5<span class="elsevierStyleHsp" style=""></span>mV. Rats with VLS lesions demonstrated the greatest amplitude in baseline activity, from 0.4 to 1.3<span class="elsevierStyleHsp" style=""></span>mV (<a class="elsevierStyleCrossRef" href="#fig0005">Fig. 1</a>B).</p><elsevierMultimedia ident="fig0005"></elsevierMultimedia><p id="par0080" class="elsevierStylePara elsevierViewall">The modal value of the amplitude in temporal muscle baseline activity differed for each of the 3 groups. The mode was 0.2<span class="elsevierStyleHsp" style=""></span>mV for control group rats, 0.4<span class="elsevierStyleHsp" style=""></span>mV for haloperidol group rats, and 0.5<span class="elsevierStyleHsp" style=""></span>mV for rats in the lesion group (<a class="elsevierStyleCrossRef" href="#fig0005">Fig. 1</a>).</p></span><span id="sec0060" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0120">Electromyographic comparison of baseline activity</span><p id="par0085" class="elsevierStylePara elsevierViewall">We found differences in mean amplitude of baseline EMG activity upon comparing the 3 groups (<span class="elsevierStyleItalic">χ</span><span class="elsevierStyleSup">2</span><span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>6.6, <span class="elsevierStyleItalic">P</span><span class="elsevierStyleHsp" style=""></span><<span class="elsevierStyleHsp" style=""></span>.03). Nested factors in the model did not exert an effect (<span class="elsevierStyleItalic">P</span><span class="elsevierStyleHsp" style=""></span>><span class="elsevierStyleHsp" style=""></span>.05) given that the greatest amplitude was observed in the group with VLS lesion and not among animals treated with haloperidol. Both of these study groups contrasted with the control group, which showed less activity at 0.3<span class="elsevierStyleHsp" style=""></span>mV (<a class="elsevierStyleCrossRef" href="#fig0010">Fig. 2</a>).</p><elsevierMultimedia ident="fig0010"></elsevierMultimedia></span><span id="sec0065" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0125">Electromyographic recordings during tremulous jaw movements</span><p id="par0090" class="elsevierStylePara elsevierViewall">The VLS lesion induced TJMs in all animals as of 24<span class="elsevierStyleHsp" style=""></span>hours after the intervention; systemic exposure to 1.0<span class="elsevierStyleHsp" style=""></span>mg/kg haloperidol also elicited TJMs in all animals by day 14 of treatment.</p><p id="par0095" class="elsevierStylePara elsevierViewall">The frequency distribution for TJM varied between the groups; rats with a ventrolateral striatum lesion displayed higher EMG amplitude and frequency than rats treated with haloperidol (<a class="elsevierStyleCrossRef" href="#fig0015">Fig. 3</a>). The modal value of TJM in animals exposed to haloperidol (H) fell in the frequency category of 3<span class="elsevierStyleHsp" style=""></span>Hz and tremor was only recorded within the interval of 3 to 5<span class="elsevierStyleHsp" style=""></span>Hz (<a class="elsevierStyleCrossRef" href="#fig0015">Fig. 3</a>A and C). In contrast, rats subjected to VLS lesion displayed 7 categories of TJM frequency between 3 and 9<span class="elsevierStyleHsp" style=""></span>Hz; the mode corresponded to the 4<span class="elsevierStyleHsp" style=""></span>Hz category (<a class="elsevierStyleCrossRef" href="#fig0015">Fig. 3</a>A).</p><elsevierMultimedia ident="fig0015"></elsevierMultimedia><p id="par0100" class="elsevierStylePara elsevierViewall">EMG activity recordings taken during TJM in rats treated with haloperidol showed amplitudes between 2 and 4<span class="elsevierStyleHsp" style=""></span>mV, while rats with VLS lesion displayed amplitudes between 3 and 9<span class="elsevierStyleHsp" style=""></span>mV (<a class="elsevierStyleCrossRef" href="#fig0015">Fig. 3</a>B and C). Furthermore, modal values for TJM occurrence also differed; among animals treated with haloperidol, amplitude was 2<span class="elsevierStyleHsp" style=""></span>mV, while amplitude was recorded in the categories of 3 and 4<span class="elsevierStyleHsp" style=""></span>mV for rats with VLS lesion (<a class="elsevierStyleCrossRef" href="#fig0015">Fig. 3</a>B).</p></span><span id="sec0070" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0130">Electromyographic comparison of tremulous jaw movements</span><p id="par0105" class="elsevierStylePara elsevierViewall">Mean values for tremor amplitude (mV) varied between the haloperidol group and the VLS lesion group (<span class="elsevierStyleItalic">χ</span><span class="elsevierStyleSup">2</span><span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>10.3, <span class="elsevierStyleItalic">P</span><span class="elsevierStyleHsp" style=""></span><<span class="elsevierStyleHsp" style=""></span>.001). Nested factors in the model did not contribute to the contrast effect (<span class="elsevierStyleItalic">P</span><span class="elsevierStyleHsp" style=""></span>><span class="elsevierStyleHsp" style=""></span>.05), which shows that the highest mean amplitude was in subjects with VLS lesion and not in animals treated with haloperidol (<a class="elsevierStyleCrossRef" href="#fig0020">Fig. 4</a>A).</p><elsevierMultimedia ident="fig0020"></elsevierMultimedia><p id="par0110" class="elsevierStylePara elsevierViewall">Frequency recordings (Hz) also showed the same type of response when the two rat groups were contrasted (<span class="elsevierStyleItalic">χ</span><span class="elsevierStyleSup">2</span><span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>31, <span class="elsevierStyleItalic">P</span><span class="elsevierStyleHsp" style=""></span><<span class="elsevierStyleHsp" style=""></span>.001). In this model, the effect of individuals nested within the group provided contrast (<span class="elsevierStyleItalic">χ</span><span class="elsevierStyleSup">2</span><span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>7, <span class="elsevierStyleItalic">P</span><span class="elsevierStyleHsp" style=""></span><<span class="elsevierStyleHsp" style=""></span>.03) and pseudoreplications did not vary when nested by rat (<span class="elsevierStyleItalic">χ</span><span class="elsevierStyleSup">2</span><span class="elsevierStyleHsp" style=""></span>=<span class="elsevierStyleHsp" style=""></span>1.7, <span class="elsevierStyleItalic">P</span><span class="elsevierStyleHsp" style=""></span><<span class="elsevierStyleHsp" style=""></span>.9). A higher mean frequency value was therefore also recorded in rats with VLS lesion than in rats treated with haloperidol (<a class="elsevierStyleCrossRef" href="#fig0020">Fig. 4</a>B).</p></span></span><span id="sec0075" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0135">Discussion</span><p id="par0115" class="elsevierStylePara elsevierViewall">The EMG description of tremulous jaw movements caused by bilateral electrolytic lesion to the VLS or by subchronic treatment with IP haloperidol shows clear differences between the frequency and amplitude patterns.</p><p id="par0120" class="elsevierStylePara elsevierViewall">TJMs are generated when the ventrolateral region of the striatum suffers an electrolytic lesion. This result is similar to that recorded in a study of dopamine depletion with 6-hydroxydopamine that induced TJM<a class="elsevierStyleCrossRefs" href="#bib0030"><span class="elsevierStyleSup">6,7</span></a> and attributed motor control over the mouth and limbs to the ventrolateral striatum.<a class="elsevierStyleCrossRefs" href="#bib0040"><span class="elsevierStyleSup">8,9</span></a></p><p id="par0125" class="elsevierStylePara elsevierViewall">The VLS is the critical striatal subregion in which the mechanisms of DA and ACh receptors and adenosine interact to regulate TJMs.<a class="elsevierStyleCrossRefs" href="#bib0025"><span class="elsevierStyleSup">5,12</span></a> It is within this context that TJMs may be induced by manipulating a number of different pharmacological and neurochemical conditions, and also by means of electrolytic lesion in this region, as shown by this study.</p><p id="par0130" class="elsevierStylePara elsevierViewall">TJMs caused by lesion to the VLS behave similarly to those described by Salamone et al.<a class="elsevierStyleCrossRef" href="#bib0065"><span class="elsevierStyleSup">13</span></a> in animals that have undergone dopaminergic manipulation. Both conditions generate periodic bursts of vertical oscillations of the lower jaw without there being any particular stimulus. These oscillations are observed during the jaw-closing phase and the transition between jaw closing and opening, as described by Cousins et al.<a class="elsevierStyleCrossRef" href="#bib0070"><span class="elsevierStyleSup">14</span></a> in rats with TJM induced with IP tacrine (acetylcholinesterase inhibitor). Furthermore, EMG activation is coherent with that reported in the temporal muscle under different functional conditions, such as chewing food<a class="elsevierStyleCrossRef" href="#bib0070"><span class="elsevierStyleSup">14</span></a> or when teeth are chattering (unpublished data, Herrera-Meza).</p><p id="par0135" class="elsevierStylePara elsevierViewall">TJM induced by electrolytic lesion to the VLS shows specific frequencies and amplitudes in the EMG study. According to EMG findings from each group of rats, tremor presents within a frequency range of 3 to 9<span class="elsevierStyleHsp" style=""></span>Hz and with an amplitude of 3 to 9<span class="elsevierStyleHsp" style=""></span>mV. This jaw activity shows frequency and amplitude values that are greater than those observed in tremor caused by dopaminergic antagonism elicited by prolonged administration of IP haloperidol (3–5<span class="elsevierStyleHsp" style=""></span>Hz and 2–4<span class="elsevierStyleHsp" style=""></span>mV). The frequency pattern for jaw activity during lesion-induced TJM is consistent with EMG findings from TJM described by Salamone et al.<a class="elsevierStyleCrossRef" href="#bib0065"><span class="elsevierStyleSup">13</span></a> in the rat parkinsonian model, and with the resting tremor (3 to 7<span class="elsevierStyleHsp" style=""></span>Hz) observed in human parkinsonism.<a class="elsevierStyleCrossRefs" href="#bib0115"><span class="elsevierStyleSup">20,21</span></a> It does not resemble the frequency characteristic of tardive dyskinesia (1–2<span class="elsevierStyleHsp" style=""></span>Hz).<a class="elsevierStyleCrossRef" href="#bib0110"><span class="elsevierStyleSup">22</span></a></p><p id="par0140" class="elsevierStylePara elsevierViewall">The EMG description showed that baseline activity of motor neurons that innervate the temporal muscle intensifies in the presence of a VLS lesion or in response to subchronic treatment with haloperidol. The amplitude of baseline activity is greater in animals with a bilateral electrolytic lesion in the VLS than in rats subjected to subchronic treatment with IP haloperidol. As a result, we understand that EMG characteristics of tremors and the increase in amplitude of baseline muscle activity may be related to the type of changes occurring in the striatum. Haloperidol is a neuroleptic drug that is relatively highly selective for the D<span class="elsevierStyleInf">2</span> receptor and causes extrapyramidal effects. Dopaminergic antagonism with that drug acts on the basal ganglia, especially the striate and related nuclei, and it also probably affects striatal neurons expressing D<span class="elsevierStyleInf">2</span> receptors in the indirect pathway. The electrolytic lesion destroys striatal neurons in the ventrolateral area of the striatum. This is the main point of entry for the nigrostriatal dopaminergic pathway in which the mechanisms of DA and ACh receptors interact with adenosine,<a class="elsevierStyleCrossRefs" href="#bib0025"><span class="elsevierStyleSup">5,12,13</span></a> which may cause generalised striatal damage with an effect on related nuclei.</p><p id="par0145" class="elsevierStylePara elsevierViewall">The EMG description of tremulous jaw movements due to electrolytic VLS lesion shows that this activity displays specific frequency and amplitude traits that can be distinguished from those observed in rats on subchronic treatment with haloperidol. However, lesion-induced tremors correspond to the tremulous jaw movements that result from a number of neurochemical and pharmacological conditions, as Salamone et al. have described.<a class="elsevierStyleCrossRef" href="#bib0065"><span class="elsevierStyleSup">13</span></a></p><p id="par0150" class="elsevierStylePara elsevierViewall">In summary, both the variation in baseline activity observed in these groups and the rhythmic activation pattern of the temporal muscle during TJM, induced either by VLS lesion or haloperidol, provide evidence that striatal mechanisms are related to motor neurons that innervate the trigeminal nerve. Although few test subjects were used to provide this description, we were able to determine the response patterns for TJMs induced by VLS lesion. EMG findings for frequency and amplitude in tremulous jaw movements caused by VLS lesion differ from the frequency and amplitude measured in animals treated with haloperidol. This variability may be attributed to the type of striatal change elicited.</p></span><span id="sec0080" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0140">Fundings</span><p id="par0155" class="elsevierStylePara elsevierViewall">This study was completed as a part of GHM's doctoral thesis under <span class="elsevierStyleGrantSponsor" id="gs1">CONACyT</span> grant ID <span class="elsevierStyleGrantNumber" refid="gs1">377111</span>. It received funding from <span class="elsevierStyleGrantSponsor" id="gs2">PROMEP-México PTC-195</span> and the <span class="elsevierStyleGrantSponsor" id="gs3">Mexican Ministry of Science and Innovation</span> [<span class="elsevierStyleGrantNumber" refid="gs3">PSI2011-29181</span>].</p></span><span id="sec0085" class="elsevierStyleSection elsevierViewall"><span class="elsevierStyleSectionTitle" id="sect0145">Conflicts of interest</span><p id="par0160" class="elsevierStylePara elsevierViewall">The authors have no conflicts of interest to declare.</p></span></span>" "textoCompletoSecciones" => array:1 [ "secciones" => array:12 [ 0 => array:2 [ "identificador" => "xres370412" "titulo" => array:5 [ 0 => "Abstract" 1 => "Introduction" 2 => "Method" 3 => "Results" 4 => "Conclusions" ] ] 1 => array:2 [ "identificador" => "xpalclavsec349651" "titulo" => "Keywords" ] 2 => array:2 [ "identificador" => "xres370413" "titulo" => array:5 [ 0 => "Resumen" 1 => "Introducción" 2 => "Método" 3 => "Resultados" 4 => "Conclusiones" ] ] 3 => array:2 [ "identificador" => "xpalclavsec349650" "titulo" => "Palabras clave" ] 4 => array:2 [ "identificador" => "sec0005" "titulo" => "Introduction" ] 5 => array:3 [ "identificador" => "sec0010" "titulo" => "Materials and methods" "secciones" => array:7 [ 0 => array:2 [ "identificador" => "sec0015" "titulo" => "Experimental subjects" ] 1 => array:2 [ "identificador" => "sec0020" "titulo" => "Study groups" ] 2 => array:2 [ "identificador" => "sec0025" "titulo" => "Electrolytic lesion to the ventrolateral striatum" ] 3 => array:2 [ "identificador" => "sec0030" "titulo" => "Implanting electrodes for electromyography recordings" ] 4 => array:2 [ "identificador" => "sec0035" "titulo" => "Recording tremulous jaw movement" ] 5 => array:2 [ "identificador" => "sec0040" "titulo" => "Perfusion" ] 6 => array:2 [ "identificador" => "sec0045" "titulo" => "Statistical analysis" ] ] ] 6 => array:3 [ "identificador" => "sec0050" "titulo" => "Results" "secciones" => array:4 [ 0 => array:2 [ "identificador" => "sec0055" "titulo" => "Electromyography recordings of baseline activity" ] 1 => array:2 [ "identificador" => "sec0060" "titulo" => "Electromyographic comparison of baseline activity" ] 2 => array:2 [ "identificador" => "sec0065" "titulo" => "Electromyographic recordings during tremulous jaw movements" ] 3 => array:2 [ "identificador" => "sec0070" "titulo" => "Electromyographic comparison of tremulous jaw movements" ] ] ] 7 => array:2 [ "identificador" => "sec0075" "titulo" => "Discussion" ] 8 => array:2 [ "identificador" => "sec0080" "titulo" => "Fundings" ] 9 => array:2 [ "identificador" => "sec0085" "titulo" => "Conflicts of interest" ] 10 => array:2 [ "identificador" => "xack93085" "titulo" => "Acknowledgements" ] 11 => array:1 [ "titulo" => "References" ] ] ] "pdfFichero" => "main.pdf" "tienePdf" => true "fechaRecibido" => "2013-10-03" "fechaAceptado" => "2013-10-13" "PalabrasClave" => array:2 [ "en" => array:1 [ 0 => array:4 [ "clase" => "keyword" "titulo" => "Keywords" "identificador" => "xpalclavsec349651" "palabras" => array:6 [ 0 => "Tremulous jaw movement" 1 => "Ventrolateral striatum" 2 => "Electrolytic lesion" 3 => "Haloperidol" 4 => "Muscle activity" 5 => "Electromyography" ] ] ] "es" => array:1 [ 0 => array:4 [ "clase" => "keyword" "titulo" => "Palabras clave" "identificador" => "xpalclavsec349650" "palabras" => array:6 [ 0 => "Temblor mandibular" 1 => "Estriado ventrolateral" 2 => "Lesión electrolítica" 3 => "Haloperidol" 4 => "Actividad muscular" 5 => "Electromiografía" ] ] ] ] "tieneResumen" => true "resumen" => array:2 [ "en" => array:2 [ "titulo" => "Abstract" "resumen" => "<span class="elsevierStyleSectionTitle" id="sect0010">Introduction</span><p id="spar0025" class="elsevierStyleSimplePara elsevierViewall">Tremulous jaw movement (TJMs) in rats can be induced pharmacologically by striatal dopaminergic manipulation or electrolytic lesion of ventrolateral striatum (VLS). This tremor has neurochemical, anatomical and electromyographic (EMG) characteristics similar to those of tremor in Parkinson patients. However, the EMG characteristics of tremors generated by electrolytic lesion to the VLS have not yet been studied.</p> <span class="elsevierStyleSectionTitle" id="sect0015">Method</span><p id="spar0030" class="elsevierStyleSimplePara elsevierViewall">This study used electromyography to describe tremulous jaw movement generated by bilateral electrolytic lesion in the VLS and compared it to tremors induced using subchronic IP treatment with haloperidol, a dopaminergic D2 receptor antagonist. The experimental groups contained rats with a lesion in the ventrolateral striatum and rats on subchronic haloperidol treatment; the control group received only the vehicle. The EMG signal from the temporal muscle was recorded at baseline and during TJMs in all groups.</p> <span class="elsevierStyleSectionTitle" id="sect0020">Results</span><p id="spar0035" class="elsevierStyleSimplePara elsevierViewall">TMJ frequencies were heterogeneous among the groups. Rats with VLS lesion showed higher amplitude and frequency values than the haloperidol-treated rats. Amplitudes at baseline also differed among the groups.</p> <span class="elsevierStyleSectionTitle" id="sect0025">Conclusions</span><p id="spar0040" class="elsevierStyleSimplePara elsevierViewall">We conclude that TMJs associated with electrolytic lesion to the VLS show a higher frequency and amplitude than tremors induced by haloperidol. This may be related to the way striatum neurons are affected.</p>" ] "es" => array:2 [ "titulo" => "Resumen" "resumen" => "<span class="elsevierStyleSectionTitle" id="sect0035">Introducción</span><p id="spar0005" class="elsevierStyleSimplePara elsevierViewall">El temblor mandibular (TM) en la rata es inducido farmacológicamente por la manipulación dopaminérgica estriatal y por lesión del estriado ventrolateral (EVL). Este temblor tiene características neuroquímicas, anatómicas y electromiográficas similares al temblor que presentan los pacientes con parkinsonismo. Pero se desconocen las características electromiográficas de los temblores generados por la lesión electrolítica del EVL.</p> <span class="elsevierStyleSectionTitle" id="sect0040">Método</span><p id="spar0010" class="elsevierStyleSimplePara elsevierViewall">En ese estudio, se describió electromiográficamente el temblor mandibular generado por la lesión electrolítica bilateral del EVL y se comparó con el inducido por el tratamiento subcrónico (i.p.) con haloperidol, neuroléptico de selectividad alta como antagonista dopaminérgico del receptor D2. A ratas con lesión en la región ventrolateral del estriado, con un tratamiento subcrónico de haloperidol, y a un grupo control que solo recibió el vehículo, se les registró la actividad electromiografía del músculo temporal en condiciones basales y durante los TM.</p> <span class="elsevierStyleSectionTitle" id="sect0045">Resultados</span><p id="spar0015" class="elsevierStyleSimplePara elsevierViewall">La distribución de frecuencias del TM entre los grupos varió, puesto que las ratas con la lesión en el EVL mostraron TM de mayor amplitud y frecuencia EMG que las ratas tratadas con el haloperidol. La amplitud en condiciones basales difirió en los distintos grupos de ratas.</p> <span class="elsevierStyleSectionTitle" id="sect0050">Conclusiones</span><p id="spar0020" class="elsevierStyleSimplePara elsevierViewall">Se concluye que los TM asociados a la lesión electrolítica del EVL son de mayor amplitud y frecuencia que los generados por haloperidol, esto puede estar relacionado con el tipo de afectación estriatal.</p>" ] ] "NotaPie" => array:1 [ 0 => array:2 [ "etiqueta" => "☆" "nota" => "<p class="elsevierStyleNotepara" id="npar0005">Please cite this article as: Herrera-Meza G, Manzo J, Hernández ME, Miquel M, García LI. Inducción del temblor mandibular por lesión electrolítica del estriado ventrolateral y por el tratamiento subcrónico con haloperidol en rata macho: un contraste electromiográfico. Neurología. 2014;29:416–422.</p>" ] ] "multimedia" => array:5 [ 0 => array:7 [ "identificador" => "fig0005" "etiqueta" => "Figure 1" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr1.jpeg" "Alto" => 1997 "Ancho" => 1346 "Tamanyo" => 111263 ] ] "descripcion" => array:1 [ "en" => "<p id="spar0045" class="elsevierStyleSimplePara elsevierViewall">(A) One-second EMG trace showing baseline activity in the temporal muscle for control group (C), haloperidol group (H), and electrolytic VLS lesion (L). Horizontal calibration: 200<span class="elsevierStyleHsp" style=""></span>ms. Vertical calibration: 1<span class="elsevierStyleHsp" style=""></span>mV. (B) Amplitude of baseline activity (mV) during the EMG recording in the temporal muscle in the control group (C), the haloperidol group (H), and the electrolytic VLS lesion group (L).</p>" ] ] 1 => array:7 [ "identificador" => "fig0010" "etiqueta" => "Figure 2" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr2.jpeg" "Alto" => 975 "Ancho" => 1360 "Tamanyo" => 29564 ] ] "descripcion" => array:1 [ "en" => "<p id="spar0050" class="elsevierStyleSimplePara elsevierViewall">Mean value<span class="elsevierStyleHsp" style=""></span>±<span class="elsevierStyleHsp" style=""></span>SD of the amplitude (mV) of EMG activity in the temporal muscle at baseline in the control group, haloperidol group, and electrolytic VLS lesion group.</p>" ] ] 2 => array:7 [ "identificador" => "fig0015" "etiqueta" => "Figure 3" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr3.jpeg" "Alto" => 2716 "Ancho" => 2171 "Tamanyo" => 209525 ] ] "descripcion" => array:1 [ "en" => "<p id="spar0055" class="elsevierStyleSimplePara elsevierViewall">(A) Frequency (Hz) of EMG activity in the temporal muscle during tremulous jaw movement in the haloperidol group (H) and the electrolytic VLS lesion group (L). (B) Amplitude (mV) of EMG activity in the temporal muscle during tremulous jaw movement in the haloperidol group (H) and the electrolytic VLS lesion group (L). (C) One-second sample of EMG trace showing tremulous jaw movements in the haloperidol group (H), and electrolytic VLS lesion group (L). Horizontal calibration: 200<span class="elsevierStyleHsp" style=""></span>ms. Vertical calibration: 1<span class="elsevierStyleHsp" style=""></span>mV.</p>" ] ] 3 => array:7 [ "identificador" => "fig0020" "etiqueta" => "Figure 4" "tipo" => "MULTIMEDIAFIGURA" "mostrarFloat" => true "mostrarDisplay" => false "figura" => array:1 [ 0 => array:4 [ "imagen" => "gr4.jpeg" "Alto" => 2088 "Ancho" => 1008 "Tamanyo" => 44199 ] ] "descripcion" => array:1 [ "en" => "<p id="spar0060" class="elsevierStyleSimplePara elsevierViewall">Mean value<span class="elsevierStyleHsp" style=""></span>±<span class="elsevierStyleHsp" style=""></span>SD of the amplitude (mV) and frequency (Hz) of EMG activity in the temporal muscle during tremulous jaw movement in rats subjected to haloperidol exposure or to electrolytic lesion in the VLS.</p>" ] ] 4 => array:5 [ "identificador" => "eq0005" "tipo" => "MULTIMEDIAFORMULA" "mostrarFloat" => false "mostrarDisplay" => true "Formula" => array:5 [ "Matematica" => "y=G+R[G]+Ps[R]+error" "Fichero" => "si1.jpeg" "Tamanyo" => 1447 "Alto" => 15 "Ancho" => 177 ] ] ] "bibliografia" => array:2 [ "titulo" => "References" "seccion" => array:1 [ 0 => array:2 [ "identificador" => "bibs0005" "bibliografiaReferencia" => array:22 [ 0 => array:3 [ "identificador" => "bib0005" "etiqueta" => "1" "referencia" => array:1 [ 0 => array:2 [ "contribucion" => array:1 [ 0 => array:2 [ "titulo" => "Tremulous jaw movements produced by acute tacrine administration: possible relation to parkinsonian side effects" "autores" => array:1 [ 0 => array:2 [ "etal" => false "autores" => array:5 [ 0 => "A.J. 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| 2019 August | 15 | 3 | 18 |
| 2019 July | 7 | 14 | 21 |
| 2019 June | 29 | 35 | 64 |
| 2019 May | 108 | 84 | 192 |
| 2019 April | 52 | 44 | 96 |
| 2019 March | 12 | 6 | 18 |
| 2019 February | 9 | 8 | 17 |
| 2019 January | 3 | 2 | 5 |
| 2018 December | 4 | 7 | 11 |
| 2018 November | 13 | 1 | 14 |
| 2018 October | 12 | 7 | 19 |
| 2018 September | 21 | 4 | 25 |
| 2018 August | 4 | 0 | 4 |
| 2018 July | 5 | 2 | 7 |
| 2018 June | 2 | 1 | 3 |
| 2018 May | 7 | 6 | 13 |
| 2018 April | 8 | 7 | 15 |
| 2018 March | 8 | 2 | 10 |
| 2018 February | 2 | 0 | 2 |
| 2018 January | 7 | 1 | 8 |
| 2017 December | 4 | 0 | 4 |
| 2017 November | 11 | 1 | 12 |
| 2017 October | 17 | 0 | 17 |
| 2017 September | 20 | 3 | 23 |
| 2017 August | 21 | 3 | 24 |
| 2017 July | 10 | 0 | 10 |
| 2017 June | 17 | 2 | 19 |
| 2017 May | 21 | 5 | 26 |
| 2017 April | 12 | 4 | 16 |
| 2017 March | 21 | 26 | 47 |
| 2017 February | 20 | 2 | 22 |
| 2017 January | 13 | 0 | 13 |
| 2016 December | 15 | 7 | 22 |
| 2016 November | 12 | 3 | 15 |
| 2016 October | 37 | 2 | 39 |
| 2016 September | 17 | 4 | 21 |
| 2016 August | 15 | 5 | 20 |
| 2016 July | 10 | 1 | 11 |
| 2016 June | 17 | 5 | 22 |
| 2016 May | 21 | 8 | 29 |
| 2016 April | 37 | 11 | 48 |
| 2016 March | 28 | 21 | 49 |
| 2016 February | 17 | 14 | 31 |
| 2016 January | 21 | 12 | 33 |
| 2015 December | 7 | 6 | 13 |
| 2015 November | 21 | 7 | 28 |
| 2015 October | 21 | 6 | 27 |
| 2015 September | 21 | 7 | 28 |
| 2015 August | 33 | 9 | 42 |
| 2015 July | 16 | 5 | 21 |
| 2015 June | 15 | 0 | 15 |
| 2015 May | 21 | 15 | 36 |
| 2015 April | 34 | 9 | 43 |
| 2015 March | 37 | 10 | 47 |
| 2015 February | 28 | 10 | 38 |
| 2015 January | 30 | 12 | 42 |
| 2014 December | 33 | 11 | 44 |
| 2014 November | 28 | 7 | 35 |
| 2014 October | 32 | 15 | 47 |
Show all
