大鼠三叉神经脊束核和背柱核向外侧丘系核投射的神经通路研究
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摘要
躯体感觉是非听觉信号传入听觉相关核团的主要来源,这些传入信号包括本体感觉和来自躯体表面的触觉、温度觉以及痛觉。三叉神经感觉复合体和背柱核是躯体感觉系统上行传导通路中二级神经元胞体所在的部位,在接受头面部、躯干和四肢的躯体感受器的传入以及调节听觉和各种机体活动中具有重要作用。头面部的本体感觉信号和听觉信号在很多核团和脑区进行整合。目前已经确定躯体感觉信号传递至听觉系统的IC、听皮层、上橄榄复合体和蜗神经核等部位,并在这些部位进行听觉和躯体感觉的整合。
目前认为主观性耳鸣主要是由噪声或药物等损伤内耳毛细胞使其向听觉中枢的传入不平衡,引发IC等听觉中枢自发放电增加造成的。而大量的临床实践和实验研究证明,电刺激或穴位刺激躯体感受器能抑制耳鸣。所以,躯体感觉系统调节听觉系统活动,尤其是躯体感觉刺激抑制耳鸣的研究已成为热点。这些神经化学通路和生理机制的阐明对于治疗耳鸣以及耳鸣治疗方法的探索都有重大的理论意义和现实意义。
有资料显示,IC内GABA释放的减少亦会使其放电增加诱发耳鸣,而向IC内注射微量的GABA受体激动剂能显著地降低耳鸣的响度。Shore等发现电刺激三叉神经能抑制由声音诱发的IC神经元放电。因此有人推测躯体感觉传入信号通过某种神经通路增加了IC内GABA的释放,降低了IC神经元的放电,这可能是临床上刺激躯体感受器治疗耳鸣的机制之一。
在以往的研究中,重点主要放在了躯体感觉系统与听觉系统在蜗神经核和IC水平的整合关系,而对NLL与躯体感觉相互关系的研究则关注较少。NLL三个亚核内都含有GABA能神经元,且均向IC有投射,是IC内抑制传入的主要核团。电刺激DNLL能增加IC内GABA的释放,抑制由声音诱发的IC放电活动。现在认为这些抑制作用主要与声源定位有关。但是否存在Sp5、背柱核→NLL(GABA能)→IC的神经通路,经此通路将刺激躯体感受器的信号传递至NLL,进而激活NLL内向IC投射的GABA能神经元,并由此抑制IC的放电而抑制耳鸣,至今尚无详细的报道。该通路可能是躯体感觉系统影响IC神经元放电活动的重要组成部分。
有鉴于此,本文以SD大鼠为实验材料,应用神经束路追踪和免疫荧光等技术,对Sp5I和背柱核向NLL中GABA能神经元的直接投射进行了研究。实验分为三个部分:(1)Sp5I向NLL的直接定位投射研究;(2)Sp5I投射至NLL内的纤维终末与GABA能神经元之关系研究;(3)背柱核向NLL的直接定位投射研究。结果如下:
1.Sp5I向NLL的直接投射
分别将顺行追踪剂dextran-Texas Red电泳入Sp5I,逆行追踪剂FG电泳入DNLL、INLL和VNLL。观察dextran-Texas Red顺行标记纤维和终末在NLL内的分布。结果发现:①向NLL投射的神经元主要分布在较小区间内的Sp5I背侧和腹侧。②向NLL投射的纤维和终末主要分布在对侧DNLL、INLL和VNLL。③Sp5I主要向对侧下丘外侧皮质核、对侧上丘、双侧蜗神经核、对侧脑桥网状核、对侧脑桥内侧核和脑桥腹侧核等核团有投射。
2.Sp5I向NLL投射的纤维终末与GABA能神经元之关系研究
将顺行追踪剂dextran-Texas Red电泳入Sp5I,免疫荧光方法染色显示NLL内GABA能神经元。顺行标记纤维终末与GABA能神经元的重叠区域主要出现在对侧的DNLL、INLL、VNLL和下丘外侧皮质核。激光扫描共聚焦显微镜观察发现,来源于Sp5I的顺行标记纤维终末与DNLL、INLL和VNLL内的GABA能神经元有直接接触。
3.背柱核向NLL的直接投射
将顺行追踪剂dextran-Texas Red电泳入背柱核,发现顺行标记纤维和终末主要分布在对侧DNLL和VNLL,少量分布在对侧INLL。在对侧外侧丘系旁核、下丘外侧皮质核的边缘区、下橄榄核、丘间核、脑桥外侧核、脑桥内侧核和脑桥腹侧核也有顺行标记纤维和终末的分布。将逆行追踪剂FG分别电泳入DNLL、INLL和VNLL,发现逆行标记细胞主要分布在闩平面及其尾侧较小节段内的对侧薄束核和楔束核。将顺行追踪剂dextran-Texas Red电泳入背柱核,然后用免疫荧光化学反应染色显示NLL内GABA能神经元,结果未发现来源于背柱核的投射纤维终末与NLL内的GABA能神经元有接触。
这些结果提示:(1) Sp5I向NLL有直接投射,来自Sp5I的顺行标记纤维终末和NLL内的GABA能神经元有直接接触,即存在Sp5I→NLL(GABA能)→IC的神经通路;(2)背柱核向NLL有直接投射,但投射终末不与NLL内的GABA能神经元接触。这些通路的进一步确证及相应的生理功能还有待于进行超微结构和电生理学的研究。
经河北省科学技术情报研究院查新表明,本研究结果在国内外文献中尚未有相同的报道。这些成果为进一步研究刺激躯体感受器抑制耳鸣的神经机制、耳鸣形成机制以及躯体感觉信息与听觉信息在脑干听觉中枢的整合机制提供了部分形态学证据。
The somatosensory system is an important source of non-auditory inputs into various nuclei related to auditory.It coveys touch,temperature,pain from the body surface and proprioception from proprioceptive receptors of the body.The trigeminal sensory complex and the dorsal column nucleus are the sites of the secondary neurons receiving the somatosensory primary afferents from head,face,body and limbs,and regulating the acoustic activity through the efferent projections.Somatosensory and auditory signals are integrated by auditory neurons in different auditory nuclei.It has been established that somatosensory signals input to auditory neurons in the inferior colliculus(IC),auditory cortex(AC),superior olive complex(SOC) and cochlear nucleus(CN).
As recently studies supported,tinnitus arises after the damage of the inner ear hair cells by noise exposure or application of drugs.These imbalance inputs are transferred from the cochlea to auditory nucleus,and then the hyperactivity of neurons were induced in the IC or other acoustic nuclei.Clinical evidence and animal experiments demonstrated that tinnitus can be suppressed by somatosensory electrical stimulation or acupuncture point stimulation. Therefore,it has become a focus on the relationship of the somatosensory system in modulating auditory activity,especially the mechanism of tinnitus suppressed by somatosensory stimulation.These illuminations of neurochemical pathways and physiological mechanisms have important academic and practical significance which to suppress tinnitus, and to further develop treatment for tinnitus.
Several lines of evidence suggest that the increase in firing rate of neurons by reducing of GABA-release in IC induced tinnitus.The loudness of tinnitus could be markedly suppressed by microinjection of GABA receptor agonist into IC.In 2006,Jain and Shore found that electrical stimulation of the trigeminal nerve produced suppression of sound-evoked responses in IC.Thus,there is an assumption whether its somatosensory inputs increased GABA-release within IC by certain neural pathway,and the discharge of neurons will be inhibited in the IC.It may be one of the mechanisms underlying of alleviation for tinnitus by somatosensory stimulation.
Up to now,numerous physiological and anatomical studies have been conducted on the integration between somatosensory and auditory in the level of CN and IC.However,it still remains unclear that what is the relationship between the somatosensory and auditory in the level of nucleus of the lateral lemniscus(NLL).Immunocytochemical studies have shown that three subnucleus of the lateral lemniscus contain GABAergic neurons,respectively.In addition,a GABAergic projection to the IC from the NLL has been confirmed,which is known to be a major source of inhibitory inputs to the IC.Previous studies have demonstrated that the effects of electrical stimulation within the DNLL can increase GABA-release in IC, and change the discharge characteristics of IC neuron.It is considered that the roles of inhibitory inputs were related to shaping binaural and monaural responses,and to pursuing sound source.It is unknown;however,whether there are neural pathways from Sp5,dorsal column nucleus to IC relayed by GABAergic neurons in NLL,by which conveyed the somatosensory signal to NLL and the GABAergic neurons projected to the IC were activated. The neural pathway may play an important role in effects of somatosensory signals on the discharge of IC neuron.
Based on those,the following experiments were processed by using anterograde and retrograde neural tracing technique,combined with immunofluorescence in Sprague-Dawley rat.The direct projection from Sp5I and dorsal column nucleus to GABAergic neurons in NLL was studied in the following experiments.Those included three parts:(1) whether there were the direct projections from the Sp5I to the NLL;(2) whether there were the terminal endings from the Sp5I making contacts with GABAergic neurons in NLL;(3) whether there were direct projections from the dorsal column nucleus to the NLL.The results were as followed:
1.The direct projection from the Sp5I to the NLL
With the respective injection of anterograde trace dextran-Texas Red into the Sp5I and retrograde tracer FG into DNLL,1NLL and VNLL,the distribution of dextran-Texas Red-labeled fibers and terminals were observed in the NLL,and the FG-labeled neurons were observed in the Sp5I.The following were the results:①The FG-labeled neurons were located in dorsal,ventral and ventrolateral of contralateral the Sp5I.②The fibers and terminals of the Sp5I-NLL projection were located in the DNLL,INLL and VNLL.③The anterogradely labeled fibers and terminals were observed in contralaterally external cortex of the inferior colliculus,superior colliculus,pontine reticular nucleus,lateral pontine nucleus,medial pontine nucleus and bilateral CN.
2.Relationship between fibers projected from the Sp5I and GABAergic neurons in the NLL
After anterograde tracer dextran-Texas Red was injected into the Sp5I,and then frozen sections of rat's brainstem were immunostained by fluorescence with glutamic acid decarboxylase(GAD) antibody to reveal the GABAergic neurons in the NLL.The overlap distribution of anterograde labeled fibers and terminals with GABAergic neurons were found in the contralaterally DNLL,INLL,VNLL and external cortex of the inferior colliculus.The direct contacts between the dextran-Texas Red-labeled Sp5I terminal endings and GABAergic neurons in the DNLL,INLL and VNLL were found under the laser scanning confocal microscope(LSCM).
3.The direct projection from the dorsal column nucleus to the NLL
Following dextran-Texas Red was injected into the dorsal column nucleus,the anterograde labeled fibers and terminals were seen in contralaterally DNLL,INLL and VNLL. Moderate terminals were observed in paralemniscal nucleus,lateral border region of the IC, inferior olive,intercollicular nucleus,lateral pontine nucleus and ventral pontine nucleus. After FG was injected into the DNLL,INLL and VNLL,the FG-retrograde labeled neurons were found in contralateral gracile nucleus and cuneate nucleus.After anterograde trace dextran-Texas Red was injected into the dorsal column nucleus,and the GABAergic neurons were visualized by immunofluorescence,the direct contacts between the anterograde terminals and GABAergic neurons were not observed by LSCM.
The above results show that:(1) There were direct projections from the Sp5I and the dorsal column nucleus to the NLL;and the terminal endings from the Sp5I made contacts with GABAergic neurons within the DNLL,INLL and VNLL,the possible neural pathway was Sp5→NLL(GABAergic)→IC.(2) The dorsal column nucleus was found to project directly to the NLL;and there was not any contact between terminal from the Sp5I and GABAergic neuron in the NLL.To confirm this possible pathway and physiological function, further researches are still needed by electrophysiology and immunoelectron microscopy.
The results of present study were not the same report in the world ground on the indagator of Hebei Science and Technology Information Institute recently.The results provided morphological foundation for investigating the integrated mechanism of somatosensory with auditory in auditory nuclei and modulation mechanism of suppressing tinnitus by somatosensory stimulation in future.
目前认为主观性耳鸣主要是由噪声或药物等损伤内耳毛细胞使其向听觉中枢的传入不平衡,引发IC等听觉中枢自发放电增加造成的。而大量的临床实践和实验研究证明,电刺激或穴位刺激躯体感受器能抑制耳鸣。所以,躯体感觉系统调节听觉系统活动,尤其是躯体感觉刺激抑制耳鸣的研究已成为热点。这些神经化学通路和生理机制的阐明对于治疗耳鸣以及耳鸣治疗方法的探索都有重大的理论意义和现实意义。
有资料显示,IC内GABA释放的减少亦会使其放电增加诱发耳鸣,而向IC内注射微量的GABA受体激动剂能显著地降低耳鸣的响度。Shore等发现电刺激三叉神经能抑制由声音诱发的IC神经元放电。因此有人推测躯体感觉传入信号通过某种神经通路增加了IC内GABA的释放,降低了IC神经元的放电,这可能是临床上刺激躯体感受器治疗耳鸣的机制之一。
在以往的研究中,重点主要放在了躯体感觉系统与听觉系统在蜗神经核和IC水平的整合关系,而对NLL与躯体感觉相互关系的研究则关注较少。NLL三个亚核内都含有GABA能神经元,且均向IC有投射,是IC内抑制传入的主要核团。电刺激DNLL能增加IC内GABA的释放,抑制由声音诱发的IC放电活动。现在认为这些抑制作用主要与声源定位有关。但是否存在Sp5、背柱核→NLL(GABA能)→IC的神经通路,经此通路将刺激躯体感受器的信号传递至NLL,进而激活NLL内向IC投射的GABA能神经元,并由此抑制IC的放电而抑制耳鸣,至今尚无详细的报道。该通路可能是躯体感觉系统影响IC神经元放电活动的重要组成部分。
有鉴于此,本文以SD大鼠为实验材料,应用神经束路追踪和免疫荧光等技术,对Sp5I和背柱核向NLL中GABA能神经元的直接投射进行了研究。实验分为三个部分:(1)Sp5I向NLL的直接定位投射研究;(2)Sp5I投射至NLL内的纤维终末与GABA能神经元之关系研究;(3)背柱核向NLL的直接定位投射研究。结果如下:
1.Sp5I向NLL的直接投射
分别将顺行追踪剂dextran-Texas Red电泳入Sp5I,逆行追踪剂FG电泳入DNLL、INLL和VNLL。观察dextran-Texas Red顺行标记纤维和终末在NLL内的分布。结果发现:①向NLL投射的神经元主要分布在较小区间内的Sp5I背侧和腹侧。②向NLL投射的纤维和终末主要分布在对侧DNLL、INLL和VNLL。③Sp5I主要向对侧下丘外侧皮质核、对侧上丘、双侧蜗神经核、对侧脑桥网状核、对侧脑桥内侧核和脑桥腹侧核等核团有投射。
2.Sp5I向NLL投射的纤维终末与GABA能神经元之关系研究
将顺行追踪剂dextran-Texas Red电泳入Sp5I,免疫荧光方法染色显示NLL内GABA能神经元。顺行标记纤维终末与GABA能神经元的重叠区域主要出现在对侧的DNLL、INLL、VNLL和下丘外侧皮质核。激光扫描共聚焦显微镜观察发现,来源于Sp5I的顺行标记纤维终末与DNLL、INLL和VNLL内的GABA能神经元有直接接触。
3.背柱核向NLL的直接投射
将顺行追踪剂dextran-Texas Red电泳入背柱核,发现顺行标记纤维和终末主要分布在对侧DNLL和VNLL,少量分布在对侧INLL。在对侧外侧丘系旁核、下丘外侧皮质核的边缘区、下橄榄核、丘间核、脑桥外侧核、脑桥内侧核和脑桥腹侧核也有顺行标记纤维和终末的分布。将逆行追踪剂FG分别电泳入DNLL、INLL和VNLL,发现逆行标记细胞主要分布在闩平面及其尾侧较小节段内的对侧薄束核和楔束核。将顺行追踪剂dextran-Texas Red电泳入背柱核,然后用免疫荧光化学反应染色显示NLL内GABA能神经元,结果未发现来源于背柱核的投射纤维终末与NLL内的GABA能神经元有接触。
这些结果提示:(1) Sp5I向NLL有直接投射,来自Sp5I的顺行标记纤维终末和NLL内的GABA能神经元有直接接触,即存在Sp5I→NLL(GABA能)→IC的神经通路;(2)背柱核向NLL有直接投射,但投射终末不与NLL内的GABA能神经元接触。这些通路的进一步确证及相应的生理功能还有待于进行超微结构和电生理学的研究。
经河北省科学技术情报研究院查新表明,本研究结果在国内外文献中尚未有相同的报道。这些成果为进一步研究刺激躯体感受器抑制耳鸣的神经机制、耳鸣形成机制以及躯体感觉信息与听觉信息在脑干听觉中枢的整合机制提供了部分形态学证据。
The somatosensory system is an important source of non-auditory inputs into various nuclei related to auditory.It coveys touch,temperature,pain from the body surface and proprioception from proprioceptive receptors of the body.The trigeminal sensory complex and the dorsal column nucleus are the sites of the secondary neurons receiving the somatosensory primary afferents from head,face,body and limbs,and regulating the acoustic activity through the efferent projections.Somatosensory and auditory signals are integrated by auditory neurons in different auditory nuclei.It has been established that somatosensory signals input to auditory neurons in the inferior colliculus(IC),auditory cortex(AC),superior olive complex(SOC) and cochlear nucleus(CN).
As recently studies supported,tinnitus arises after the damage of the inner ear hair cells by noise exposure or application of drugs.These imbalance inputs are transferred from the cochlea to auditory nucleus,and then the hyperactivity of neurons were induced in the IC or other acoustic nuclei.Clinical evidence and animal experiments demonstrated that tinnitus can be suppressed by somatosensory electrical stimulation or acupuncture point stimulation. Therefore,it has become a focus on the relationship of the somatosensory system in modulating auditory activity,especially the mechanism of tinnitus suppressed by somatosensory stimulation.These illuminations of neurochemical pathways and physiological mechanisms have important academic and practical significance which to suppress tinnitus, and to further develop treatment for tinnitus.
Several lines of evidence suggest that the increase in firing rate of neurons by reducing of GABA-release in IC induced tinnitus.The loudness of tinnitus could be markedly suppressed by microinjection of GABA receptor agonist into IC.In 2006,Jain and Shore found that electrical stimulation of the trigeminal nerve produced suppression of sound-evoked responses in IC.Thus,there is an assumption whether its somatosensory inputs increased GABA-release within IC by certain neural pathway,and the discharge of neurons will be inhibited in the IC.It may be one of the mechanisms underlying of alleviation for tinnitus by somatosensory stimulation.
Up to now,numerous physiological and anatomical studies have been conducted on the integration between somatosensory and auditory in the level of CN and IC.However,it still remains unclear that what is the relationship between the somatosensory and auditory in the level of nucleus of the lateral lemniscus(NLL).Immunocytochemical studies have shown that three subnucleus of the lateral lemniscus contain GABAergic neurons,respectively.In addition,a GABAergic projection to the IC from the NLL has been confirmed,which is known to be a major source of inhibitory inputs to the IC.Previous studies have demonstrated that the effects of electrical stimulation within the DNLL can increase GABA-release in IC, and change the discharge characteristics of IC neuron.It is considered that the roles of inhibitory inputs were related to shaping binaural and monaural responses,and to pursuing sound source.It is unknown;however,whether there are neural pathways from Sp5,dorsal column nucleus to IC relayed by GABAergic neurons in NLL,by which conveyed the somatosensory signal to NLL and the GABAergic neurons projected to the IC were activated. The neural pathway may play an important role in effects of somatosensory signals on the discharge of IC neuron.
Based on those,the following experiments were processed by using anterograde and retrograde neural tracing technique,combined with immunofluorescence in Sprague-Dawley rat.The direct projection from Sp5I and dorsal column nucleus to GABAergic neurons in NLL was studied in the following experiments.Those included three parts:(1) whether there were the direct projections from the Sp5I to the NLL;(2) whether there were the terminal endings from the Sp5I making contacts with GABAergic neurons in NLL;(3) whether there were direct projections from the dorsal column nucleus to the NLL.The results were as followed:
1.The direct projection from the Sp5I to the NLL
With the respective injection of anterograde trace dextran-Texas Red into the Sp5I and retrograde tracer FG into DNLL,1NLL and VNLL,the distribution of dextran-Texas Red-labeled fibers and terminals were observed in the NLL,and the FG-labeled neurons were observed in the Sp5I.The following were the results:①The FG-labeled neurons were located in dorsal,ventral and ventrolateral of contralateral the Sp5I.②The fibers and terminals of the Sp5I-NLL projection were located in the DNLL,INLL and VNLL.③The anterogradely labeled fibers and terminals were observed in contralaterally external cortex of the inferior colliculus,superior colliculus,pontine reticular nucleus,lateral pontine nucleus,medial pontine nucleus and bilateral CN.
2.Relationship between fibers projected from the Sp5I and GABAergic neurons in the NLL
After anterograde tracer dextran-Texas Red was injected into the Sp5I,and then frozen sections of rat's brainstem were immunostained by fluorescence with glutamic acid decarboxylase(GAD) antibody to reveal the GABAergic neurons in the NLL.The overlap distribution of anterograde labeled fibers and terminals with GABAergic neurons were found in the contralaterally DNLL,INLL,VNLL and external cortex of the inferior colliculus.The direct contacts between the dextran-Texas Red-labeled Sp5I terminal endings and GABAergic neurons in the DNLL,INLL and VNLL were found under the laser scanning confocal microscope(LSCM).
3.The direct projection from the dorsal column nucleus to the NLL
Following dextran-Texas Red was injected into the dorsal column nucleus,the anterograde labeled fibers and terminals were seen in contralaterally DNLL,INLL and VNLL. Moderate terminals were observed in paralemniscal nucleus,lateral border region of the IC, inferior olive,intercollicular nucleus,lateral pontine nucleus and ventral pontine nucleus. After FG was injected into the DNLL,INLL and VNLL,the FG-retrograde labeled neurons were found in contralateral gracile nucleus and cuneate nucleus.After anterograde trace dextran-Texas Red was injected into the dorsal column nucleus,and the GABAergic neurons were visualized by immunofluorescence,the direct contacts between the anterograde terminals and GABAergic neurons were not observed by LSCM.
The above results show that:(1) There were direct projections from the Sp5I and the dorsal column nucleus to the NLL;and the terminal endings from the Sp5I made contacts with GABAergic neurons within the DNLL,INLL and VNLL,the possible neural pathway was Sp5→NLL(GABAergic)→IC.(2) The dorsal column nucleus was found to project directly to the NLL;and there was not any contact between terminal from the Sp5I and GABAergic neuron in the NLL.To confirm this possible pathway and physiological function, further researches are still needed by electrophysiology and immunoelectron microscopy.
The results of present study were not the same report in the world ground on the indagator of Hebei Science and Technology Information Institute recently.The results provided morphological foundation for investigating the integrated mechanism of somatosensory with auditory in auditory nuclei and modulation mechanism of suppressing tinnitus by somatosensory stimulation in future.
引文
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