大鼠三叉神经本体觉中枢通路的形态学和电生理学研究
摘要
三叉神经本体觉的中枢通路是神经解剖学长期以来的未解之迷之一，主要是由于方法学的限制和传导本体觉的三叉神经中脑核(Vme)神经元在中枢内的分布特殊。1979年HRP跨节追踪技术的问世给这一研究提供了前提。1981年以后，Matesz、Nomura等，彭兆知等分别用此技术发现了Vme神经元中枢突投射于同侧三叉神经脊束核吻侧亚核背内侧区和邻接的网状结构(Vodm-LRF)，形成“哑铃形”的分布区。1988年王百忍和李继硕以此为线索综合运用HRP标记技术，发现了一条大鼠三叉神经本体觉中枢通路，证明此通路由Vme→Vodm-LRF→”带状区”→对侧丘脑腹后内侧核(VPM)等四级神经元组成。“带状区”又分为三叉神经感觉主核背内侧部(Vpdm)，三叉神经运动核腹侧区(AVM)和上橄榄核背侧区(ADO)。但此通路中的Vodm-LRF和“带状区”等两个中继部位，是否有独立的中继核团存在，在此中继的前、后两级神经元间是否形成突触连接，都还是神经解剖学中的空白，此外有关此通路的电生理学研究也是必不可少的。所以本论文的目的是用形态学和电生理学方法证明上述通路确实存在，以及最初对该通路的提法的正确性。整个论文可分两大部分：前四章节证明通路中各级神经元间的联系，以及以往未作观察的二、四级神经元的细胞构筑；后两章节主要是关于Vme神经元的传入投射，以及来自通路中第二级神经元---Vodm神经元的传入投射的意义。
1．溃变法和标记法相结合的电镜研究，将Ricin注入大鼠咬肌神经，HRP分别注入Vpdm和AVM。电镜观察证明，在Vodm区初级传入的溃变终末与HRP逆标的树突形成突触，突触后树突多为中间树突和近侧
A possible proprioceptive pathway in the rat was reported by Wang and LI in 1988, the pathway was considered to be composed of four orders of neurons, those were indicated to be located in the mesencephalic trigeminal nucleus (Vme), in the dorsomedial part of subnucleus oral is (Vodm) and adjacent parvocellular reticular nucleus (PCRt), in the "ribbon-shape area" and in the contralateral ventroposterior medial thalamic nucleus (VPM). The "ribbon-shape area" had three enlargements --- dorsomedial part of principal trigeminal sensory nucleus (Vpdm), area ventral to motor trigeminal nucleus (AVM) and area dorsal to superior olivery nucleuus (ADO).
This whole dissertation was aimed at the above pathway, in order to confirm the existence of this pathway, and to prove how correct their original propose was. Thus, the whole dissertation
could be divided into two parts-part one, composed of four
studies, was about the connections of the pathway; and part two,
composed of two studies, was tend to the first order neuron -
the Vme itself.
1. In order to obtain fine structural evidence for the pathway mentioned above, firstly Ricinus communis (Ricin) was injected into the masseter nerve of the rats, and after about two weeks survival HRP was then injected into the "ribbon-shape area". It was observed that electron-dense degenerated terminals synapsed on HRP labeled dendrites. The synapses identified were all the type of asymmetry. The postsynaptic dendrites were mainly primary and intermedial ones.
Secondly, kianic acid was injected into the contralateral thalamus. Both filamentous and electron-dense degenerated boutons were seen synapsing on HRP labeled either somata or dendrites in the three enlargements of the "ribbon-shape area".
Therefor, connections between first order and second order neurons in the Vodm, and between second order and third order neurons in the "ribbon-shape area" had been electron-microscopically testified.
2. It was morphologically demonstracted in the rat that Vme neurons projected to contralateral VPM through another two orders of neuron. Based upon the morphological date, the present work furthered the research by probing functional relationship between the relay neurons mentioned above. Recording micropipette was localized in the Vodm, the Vpdm and AVM, and the contalatral VPM, respectively, unite discharges were found at first. The responses with changing of discharge frequencies to lowering down the jaw, probing the masseter muscle and electric stimulating the
1．溃变法和标记法相结合的电镜研究，将Ricin注入大鼠咬肌神经，HRP分别注入Vpdm和AVM。电镜观察证明，在Vodm区初级传入的溃变终末与HRP逆标的树突形成突触，突触后树突多为中间树突和近侧
A possible proprioceptive pathway in the rat was reported by Wang and LI in 1988, the pathway was considered to be composed of four orders of neurons, those were indicated to be located in the mesencephalic trigeminal nucleus (Vme), in the dorsomedial part of subnucleus oral is (Vodm) and adjacent parvocellular reticular nucleus (PCRt), in the "ribbon-shape area" and in the contralateral ventroposterior medial thalamic nucleus (VPM). The "ribbon-shape area" had three enlargements --- dorsomedial part of principal trigeminal sensory nucleus (Vpdm), area ventral to motor trigeminal nucleus (AVM) and area dorsal to superior olivery nucleuus (ADO).
This whole dissertation was aimed at the above pathway, in order to confirm the existence of this pathway, and to prove how correct their original propose was. Thus, the whole dissertation
could be divided into two parts-part one, composed of four
studies, was about the connections of the pathway; and part two,
composed of two studies, was tend to the first order neuron -
the Vme itself.
1. In order to obtain fine structural evidence for the pathway mentioned above, firstly Ricinus communis (Ricin) was injected into the masseter nerve of the rats, and after about two weeks survival HRP was then injected into the "ribbon-shape area". It was observed that electron-dense degenerated terminals synapsed on HRP labeled dendrites. The synapses identified were all the type of asymmetry. The postsynaptic dendrites were mainly primary and intermedial ones.
Secondly, kianic acid was injected into the contralateral thalamus. Both filamentous and electron-dense degenerated boutons were seen synapsing on HRP labeled either somata or dendrites in the three enlargements of the "ribbon-shape area".
Therefor, connections between first order and second order neurons in the Vodm, and between second order and third order neurons in the "ribbon-shape area" had been electron-microscopically testified.
2. It was morphologically demonstracted in the rat that Vme neurons projected to contralateral VPM through another two orders of neuron. Based upon the morphological date, the present work furthered the research by probing functional relationship between the relay neurons mentioned above. Recording micropipette was localized in the Vodm, the Vpdm and AVM, and the contalatral VPM, respectively, unite discharges were found at first. The responses with changing of discharge frequencies to lowering down the jaw, probing the masseter muscle and electric stimulating the
引文
1. Aldes L.D. and Boone T.B., A combined flat-embedding, HRP histochemical method for correlative light and electron microscopic study of single neurons. J. Neurosci. Res.11(1984)27-34.
2. Coyle J.T. and Schwatcz R., The use of excitatory amino acids as selective neurotoxins, In Bjorklund A. and Hokfelt T. (Eds),Handbook of Chemical Neuroanatomy, Vol. 1. Amsterdam: Elsevier Biocedical Press, 1983, pp.508-527.
3. Coyle J.Y., Molliver M.E. and Kuhar M.J., In the situ injection of kainic acid: A new method for selectively lesioning neuronal cell bodies while sparing axons of passage. J. Comp.Neurol. 180(1978)301-324.
4. Gu Y.M., Chen Y.C. and Ye L.M., A new highly sensitive HRP-TMB method using sodium tungstate as a stabilizer: II. Electron microscopic study. Chinese J. Neuroanat. 7(1991)124-129.
5. Lindsley D.F. and Holmes J.E., Basic Human Neurophysiology, Elsevier, New York/Amsterdam/Oxford, 1984, pp62-82.
6. Luo P.F., Wang B.P., Peng Z.Z. and Li J.S., Morphological characteristics and terminating patterns of masseteric neurons of the mesencephalic trigeminal nucleus in the rat: An intracellular horseradish peroxidase labeling study. J. Comp. Neurol.303(1991)286-299.
7. Moriizumi T., Nakamura Y., Okoyama S. and Kitao y., Synaptic organization of the cat entopeduncullar nucleus with special reference to the relationship between the afferents to entopedunculothalamic projection neurons: An electron microscope study by a combined degeneration and horseradish peroxidase tracing technique. Neuroscience 20(1987)797-816.
8. Mugnani E. and Friedrich V.L., Electron microscopy:Identification study of normal and degenerating neuronal elements by electron microscopy. In Heimer L. and Robards M. J. [eds.],Neuroanatomical tract-tracing methods, Plenum Press, New York and London. 1981, pp377-406.
9. Streit W.J. and Kreutzberg G.W., Response of endogenous glial cells to motor neuron degeneration induced by toxic ricin. J.Comp. Neurol. 268(1988)248-263.
10.Takeuchi Y., McLean J.H. and Hopkins D.A., Reciprocal connections between the amygdala and parabrachial nuclei: ultrastructural demonstration by degeneration and axonal transport of horseradish peroxidase in the cat. Brain Research 239(1982)583-588.
11.Wang B.R. and Li J.S., The projection pathway from the trigeminal mesencephalic nucleus (Vme) to the thalamus in the rat. I. The brain stem distribution of the "central" processes of Vme--- A light and electron microscope study. Chinese J.Neuroanat. 4(1988)35-40.
12.Wang B.R. and Li J.S., The projection pathway from the trigeminal mesencephalic nucleus (Vme) to the thalamus in the rat. II. The efferent projection of the dorsal-medial portion of the subnucleus oralis of the trigeminal spinal nucleus(Vodm) and the adjacent lateral reticular formation (LRF). Chinese J.Neuroanat. 4(1988)41-51.
13.Wang B.R. and Li J.S. , The projection pathway from the trigeminal mesencephalic nucleus (Vme) to the thalamus in the rat. III. The projection from the zone-shaped area along the medial border of principal trigeminal nucleus to the thalamus. Chinese J. Neuroanat . 4(1988)53-60.
14.Wiley R.G., Blessing W.W. and Reis D.J., Suicide transport:Destruction of neurons by retrograde transport of ricin, abrin,and modecein. Science 216(1982)889-890.
15.Xie W., Zhang J.D., Yang S. and Li J.S., Relationship between degeneration of mesencephalic trigeminal neurons induced by ricin and different amount of ricin used or different animal survival times. submitted
16.Yamamoto T., Iwasaki Y., Konno H. and kudo H., Retrograde axoplasmic transport of toxic lectins is useful for transganglionic tracings of the peripheral nerve. Brain Research 274(1983)325-328.
17.Zhang J.H., Li H.M. and Li J.S., The relationship between sensitivity, preservation of tissue ultrastructure and the pH of acetate buffer in HRP-TMB reaction medium. Chinese J. Neuroanat.3(1987)118-122.
18.Zhang J.D. and Li J.S., A possible trigeminal proprioceptive pathway from the masseter muscle to the contralateral thalamus---Comprehensive tracing studies in the rat. submitted
19.Zhang J.D. and Li J.S., Nucleus morphology, location,cytoarchitecture and connections of the secondary neurons in the trigeminal proprioceptive pathway in the rat---A light microscopic observation. Chinese J. Neuroanat. 8(1992)63-72.
20.Zhang J.D., Wang B.R., Li H.M. and Li J.S., Projections from neurons innervating the masseter muscle to the subnucleus oralis of the spinal trigeminal nucleus and adjacent lateral reticular formation in the rat. J. Hirnforsch. 32(1991)641-646.
[1] Abe K. A study of sensory projection from jaw muscles to the cerebralcortex inthe rat. Jap J Physiol. 1978;28.309～322
[2] 王百忍，李继硕．大鼠三叉神经中脑核神经元中枢突在脑干内投射的光、电镜观察(中脑核——丘脑投射通路研究之一)．神经解剖学杂志，1988；4：35～40
[3] 王百忍，李继硕．大鼠三叉神经脊束核吻侧亚核背内侧部及邻接的外侧网状结构的传出投射——WGA—HRP顺行追踪研究(中脑核——丘脑投射通路研究之二)．神经解剖学杂志，1988;4，41～52
[4] 王百忍，李继硕．大鼠沿三叉神经感觉主核内缘的带状区向丘脑的投射——HRP法研究(中脑核——丘脑投射通路研究之三)．神经解剖学杂志，1988；4：53～60
[5] Appenteng BK, Donga R, Willians RG. Morphological and electrophysiologicaldetermination of the projections of jaw--elevatormuscle spindle afferents in rats. J Physiol, 1985;369:93～113
[6] Matesz C. Peripheral and cental distribution of fibres of the mesencephalic trigeminal root in the rat. Neurosci Lett, 1981;27:13～17
[7] 罗丕福，李继硕．GABA样神经元和GABA能三叉一丘脑投射神经元在三叉神经感觉核簇的分布及形态学特征．神经解剖学杂志，1991;7：185～194
[8] Rustioni A. Non --primary afferents to the nucleus gracilis fromlumbar cord of the cat. Brain Res, 1973;51:81～95
[9] 吕国蔚．背索突触后神经元．生理科学进展，1984;15：310～315
[10] 罗丕福，李继硕．大鼠三叉神经本体觉三级传导通路的形态学证据——Ricin跨节溃变与HRP逆行追踪结合的光、电镜研究．神经解剖学杂志，1992;8：57～62
[11] Luo PF, Wang BR, Peng ZZ, et al. Morphological characteristics and terminating patterns of masseteric neurons of the mesencephalic trigeminal nucleus in the rat: An intracellular horseradish peroxidase labeling study, j Comp Neurol, 1990(303:286～299
[12] 张敬东，李继硕．大鼠三叉神经本体觉中枢通路二级神经元中继咬肌本体觉传入的超微结构证据．神经解剖学杂志，1992；8：203～210
[13] 李金莲，张敬东，李继硕．三叉神经本体觉中枢通路二、三级神经元之间突触连接的发现及突触构筑学研究——溃变与HRP逆行追踪法相结合的电镜观察．神经解剖学杂志，1993；9：97～104
1．王百忍 李继硕；神经解剖学杂志 4：35，1988
2．王百忍 李继硕：神经解剖学杂志 4：41，1988
3．王百忍 李继硕：神经解剖学杂志 4：53，1988
4．李惠民 李继硕：解剖学报 15：366，1984
5．罗丕福 李继硕：神经解剖学杂志 8：57，1992
6．彭兆知 张凤鸣：解剖学杂志 19(增)：279，1986
7．彭兆知等：神经解剖学杂志 2：27，1986
8. Alvarado-Mallart RM et al, Exp Brain Res 23:167, 1975
9. Appenteng K et al, J Physiol 369:93, 1985
10. Capra NF and Wax TD, J Comp Neurol 279:341, 1989
11. Carpenter MB and Sutin J. Human Neuroanatomy Williams & Wilkins London p 393, 1983
12. Darian-Smith I. Handbook of sensory physiology Springer-Verlag Berlin Heidelb erg New York p27l, 1973
13. Denng-Brian D and Yanagisawa N, Brain 96:7.83, 1973
14. Erzurumlu R and Killackey HP. J Comp Neurol 188:75, 1979
15. Falls WM: Brain Res 290:136, 1981
16. Falls WM, Neuroscience 13:1279, 1984
17. Falls WM et al. Somatosens Res 3:89, 1985
18. Gobel S and Purvis MB. Brain. Res 48:27, 1972
19. Huerta MF et al: J Comp Neurol 220:147, 1983
20. Kelly JP.. Principles of neural science Elsevier New York p562, 1985
21. Kruger L et al: Pain in the trigeminal rigion Elsevier New York pl91, 1977
22. Kruger L et al.. The cranial nerves Springer-Verlag Berlin p273, 1981
23. Luo PF et al, J Comp Neurol 303:286, 1991
24. Matesz C: Neurosci Lett 27:13, 1981
25. Mesulam M-M: Tracing neural connections with horseradish peroxidase John Wiley & Sons New York p 1 , 1982
26. Mizuno N et al. J Comp Neurol 215:290, 1983
27. Nishimori T et al. Brain Res 372:375, 1986
28. Nomura S and Mizuno N. Neurosci Lett Suppl 17:sl52, 198.)
29. Ruggireo DA et al: Brain Res 225:225, 1981
30. Schvarcz JR: Advances in pain research and therapy Raven. Press New York p 331, 1979
31. Shigenaga Y et al: J Comp Neurol 243:388, 1986
32. Shigenaga Y et al. J Comp Neurol 244:1, 1986
33. Shigenaga Y et al: J Comp Neurol 268:489, 1988
34. Zhang JD et al. J Hirnforsch 32:641, 1991
1．王百忍，李继硕．大鼠三叉神经中脑核神经元中枢突在脑干内投射的光、电镜观察(中脑核——丘脑投射通路研究之一)。神经解剖学杂志，1988；4：35-40
2．王百忍，李继硕．大鼠三叉神经脊束核吻侧亚核背内侧部及邻近的外侧网状结构的传出投射——WGA—HRP顺行追踪研究(中脑核——丘脑投射通路研究之二)。神经解剖学杂志，1988；4：41-52
3．王百忍，李继硕．大鼠沿三叉神经感觉主核内缘的带状区向丘脑的投射——HRP法研究(中脑核——丘脑投射通路研究之三)，神经解剖学杂志，1988；4：53～60
4．顾耀铭，陈以慈，叶鹿鸣．钨酸钠作为稳定剂的新的高更敏HRP-TMB法——Ⅱ。电镜研究。神经解剖学杂志，1991；7：124-129
5. Mesulum MM. Tetramethyl benzidine for horseradish peroxidase neuro-histochemistry. A non-carcinogenic blue reaction product with superior sensitivity for visualizing neural afferents and efferents. J Histochem Cytochem, 1978; 26:106-117
6．张敬东，李继硕．三叉神经本体觉中枢通路二级神经元所在地(三叉神经脊束核吻侧亚核背内侧区，Vodm)的核团形态、细胞构筑及纤维联系——光镜观察．神经解剖学杂志，1992；8：633-72
7. Faxinos G, Watson C. The rat. brain in stereotaxic coordinates, 2nd ed. Academic Press, Sydney/Orlando/San Diego/NewYork/Austin/London/Montreal/Tokyo/Toronto, 1986.
8．王振福，………李继硕．大鼠沿三叉神经感觉主核内缘的“带状区”向丘脑的投射和细胞构筑-HRP法研究．神经解剖学杂志，1995(在印刷中)
9. Carpenter MB, Sutin J. Human Neuroanatomy, 8th ed. Baltimore/London, Williams & Wilkins, 1983, pp.516-524
10. Beckstead RM, Morse JR, Norgren R. The nucleus of the solitary tractin the monkey:Projections to the thalamus and brain stem nuclei. J Comp Neurol, 1980; 190:259-282
11. Faull RLM, Mehler WR. Thalamus. In G Paxinos (ed.), The rat nervous system. Vol 1,Academic Press, Sydney, 1985, pp. 129-168
12. Smith RL. The ascending fiber projections from the principal sensory trigeminal nucleus in the rat. J Comp Neurol, 1973; 148:423-446
13. Smith RL. Axonal projections and connections of the principal sensory trigeminal nucleus in the monkey. J Comp Neurol, 1975; 163:347-376
14. Fukushima I, Kerr FWL. Organization of trigeminothalamic tracts and other thalamic afferent systems of the brainstem in the rat: Presence of gelatinosa neurons with thatamic connections. J Comp Neurol, 1979; 183:169-184
15. Eisenman J, Fromm G, Landgren S, et al. The ascending projections of trigeminal neurones in the cat, investigated by antidromic stimulation. Acta Physiol Scand, 1964; 60:337-350
16. Jerge CR. The organization and function of the trigeminal mesencephalic nucleus. J Neurophysiol, 1963; 26:379-392
17. Lund JP, Sessle BJ. Oral-facial and jaw muscle afferent projections to neurons in cat frontal cortex. Exp Neurol, 1974; 45:314-331
18. Abe K. A study of sensory projection from jaw muscles to the cerebral cortex in the rat. Jap J Physiol, 1978; 28:309-322
[1] Matesz C. Peripheral and central distribution of fibres of the mescnccphalic trigcminal root in the rat. Ncurosci Lett, 1981;27:13～17
[2] 彭兆知，曹广义，李继硕．大鼠三叉神经中脑核神经元中枢突的行径和分布——HRP跨节追踪研究．神经解剖学杂志，1986;2：27～36
[3] Juch PJW, Roxk JTM. Efferent projections of the parvocellular reticular nucleus to the mesencephalic trigeminal nucleus in rat. Brain Res, 1988;462:185～189
[4] Roxk JTM, Juch PJW, van Willigen JD. Arrangement and connections of mesencephalic trigeminal neurons in the rat. Acta Anat, 1986;127:7～15
[5] Zhang JD, Wang BR, Li HM, et al. Projections from neurons innervating the masscter muscle to the subnuclcus oralis of the spinal trigeminal nucleus and adjacent lateral reticular formation in the rat. J Hirnforsch, 1991;32:641~646
[6] 王百忍，李继硕．大鼠三叉神经中脑核神经元中枢突在脑干内投射的光、电镜观察(中脑核——丘脑投射通路研究之一)．神经解剖学杂志，1988;4：35～40
[7] 王百忍，李继硕．大鼠三叉神经脊束核吻侧亚核背内侧部及邻接的外侧网状结构的传出投射——WGA-HRP顺行追踪研究(中脑核——丘脑投射通路研究之二)．神经解剖学杂志，1988；4：41～52
[8] 王百忍，李继硕．大鼠沿三叉神经感觉主核内缘的带状区向丘脑的投射——HRP法研究(中脑核——丘脑投射通路研究之三)．神经解剖学杂志，1988；4：53～60
[9] 张敬东，李继硕．大鼠三叉神经本体觉中枢通路二级神经元中继咬肌本体觉传入的超微结构证据．神经解剖学杂志，1992；8：203～210
[10] 李金莲，张敬东，李继硕．三叉神经本体觉中枢通路二、三级神经元之间突触连接的发现及突触构筑学研究——溃变与HRP逆行追踪法相结合的电镜观察．神经解剖学杂志，1993;9：97～104
[11] 张敬东，李继硕．三叉神经本体觉中枢通路二级神经元所在地(三叉神经脊束核吻侧亚核背内侧区，Vodm)的核团形态、细胞构筑及纤维联系——光镜观察神经解剖学杂志，1992;8：63～72
[12] 顾耀铭，陈以慈．叶鹿鸣．钨酸钠作为稳定剂的新的高灵敏HRP-TMB法——Ⅱ、电镜研究．神经解剖学杂志，1991；7：124～129
[13] Roxk JTM, Luiten PGM, van Willigen JD. Afferent projections to the mesencephalic trigeminal nucleus in the rat. Acta Anat, 1988;132:269～274
[14] Hinrichsen CFL, Larramendi LMH?Synapses and cluster formation in the mouse mesencephalic fifth nucleus. Brain Res,1968;7:296～299
[15] Hinrichsen CFL, Larramendi LMH. The trigeminal mesencephalic nucleus IⅠ. Electron microscopy. Am J Anat, 1970;127:303～320
[16] Liem RSB, Copray JCVM , van Willigen JD. Ultrastructure of the rat mesencephalic trigeminal nucleus. Acta Anat, 1991;140:112～119
[17] Inagaki N, Yamatodani A, Shinoda K, et al. The histaminergic innervation of mesencephalic neurons of the trigeminal nerve in rat brain: a light and electron microscopical study. Brain Res, 1987;418:388～391
[18] Copray JCVM, Liem RSB, Ter Horst GJ, et al. Dopaminergic afferents to the mesencephalic trigeminal nucleus: a light and electron microscope immunocytochemistry study. Brain Res, 1990;541:343～348
[19] Copray JCVM, Ter Horst GJ, Liem RSB, et al. Neurotransmitters and neuropeptides within the mesencephalic trigeminal nucleus of the rat: An immunohistochemical analysis. Neuroscience, 1990;37:399～411
[20] 史中立，秦秉志，李继硕．大鼠骶髓后连合核的中枢联系及机能分析——HRP法研究．解剖学报，1990;21：360～368
[21] 贾宏阁，饶志仁，施际武．大鼠臂旁核内接受孤束核内脏传入之神经元同时接受中央杏仁核的反馈调节——溃变、HRP法结合包埋后免疫电镜研究．神经解剖学杂志，1994；10：294～298
[22] Kayahara T. Synaptic connections between spinal motoneurons and dorsal root ganglion cells in the cat. Brain Res, 1986; 376:299～309
[1] Hinrichsen CFL, Larrnmendi LMH. Synapses and cluster formation in the mouse metiencephalic fifth nucleus. Brain Res,1968;7:296～299
[2] Hinrichsen CFL, Laxramendi LMH. The trigeminal mesencephalic nucleus IⅠ . Electron microscopy, Arm J Anat,1970;127:303～320
[3] Liem RSB, Copray JCVM, van Willigen JD. Ultrastructure of the rat mesencephalic trigerminal nucleus. Acta Anat,1991;140:112～119
[4] Inagaki N, Yamatodani A, Shinoda K, et al. The histaminergic innervation of mesencephalic neurons of the trigeminal nerve in rat brain : a light and electron microscopical study. Brain Res, 1987 ; 418:388～391
[5] Copray JCVM , Liem RSB. Ter Horst GJ, et al. Dopaminergic afferents to the mesencephalic trigeminal nucleus; a light and electron microscope immunocytochemistry study. Brain Res. 1990 ; 514 : 343～348
[6] Copray JCVM. Ter Horst GJ, Liem RSB. et al. Neurotransmitters and neuropeptides within the mesencephalic trigeminal nucleus of the rat; an immunohistochemical analysis. Neuroscience. 1990;37: 399～411
[7] Carpenter MB, Sutin J. Hur an neuroanatomy. Baltimore'London : Williams & Wilkins, 1983:393～401
[8] Kayahara T, Takimoto T, Sakashita S. Synaptic junctions in the cat spinal ganglion. Brain Res, 1981; 216:277～290
[9] Kayahara T, Sakashita S, Takimoto T. Evidence for spinal origin of neurons synapsing with dorsal root ganglion cells of the cat. Brain Res. 1984;293:225～230
[10] Kayahara T, Synaptic connections between spinal motonenrons and dorsal root ganglion cells in the cat. Brain Res, 1986;376:299～309
[11] 李云庆，饶志仁，施际武．大鼠三叉神经感觉核簇内SP受体及三叉神经节内SP样阳性终末和SP受体分布的免疫组织化学研究．神经解剖学杂志，1993；9：89～92
[12] Peters A. Palay SL, de E Webster H. The fine structure of the nervous system. London: WB Saunders Company. 1976;118～180
[13] De Montigny C, Lund JP. A microiontophoretic study of the action of kainic acid and putative neurotransmitters in the rat Tnesenvephalic trigeminal nucleus. Neuroscience , 1980;5 ,1621～1628
[14] Guyenet PG, Aghajanian GK. Excitation of neurons in the locus coeruleus by substance P and related peptide. Brain Res, 1977;136:178～184
[15] Henderson G, Pepper CM, Shefner SA. Electrophysiological properties of neurons contained in the locus coeruleus and mesencephalic nucleus of the trigeminal nerve in vitro. Exp Brain Res, 1982;45;29～37
[16] Minkels RF, Juch PJW, Ter Horst GJ, et al. Projections of the parvocellular reticular formation to the contralaterai mesencephalic trigeminal nucleus in the rat. Brain Res, 1991;547:13～21
[17] Liem RSB, Copray JCVM, van Willigen JD. Distribution of synaptic boutons in the mesencephalic trigeminal nucleus of the rat -- A quantitative electron-microscopical study. Acta Anat, 1992; 143; 74～78
[18] Luo PF. Wang BR, Peng ZZ. et al. Morphological characteristics and terminating patterns of masseteric neurons of the mesencephalic trigeminal nucleus in the rat: An intracellular horseradish peroxidase labeling study. J Comp Neurol, 1991 ; 303:286～299
[19] Shigenaga Y. Mitsuhiro Y, Yoshida A. et al. Morphology of single mesencephalic trigeminial neurons innervating masseter muscle of the cat. Brain Res, 1988;445:392～399
[20] Shigenaga Y, Mitsuhiro Y, Shirana Y , et al. Two types of jaw-muscle spindle afferents in the cat as demonstrated by intraaxonal staining with HRP. Brain Res. 1990;514: 219～237
2. Coyle J.T. and Schwatcz R., The use of excitatory amino acids as selective neurotoxins, In Bjorklund A. and Hokfelt T. (Eds),Handbook of Chemical Neuroanatomy, Vol. 1. Amsterdam: Elsevier Biocedical Press, 1983, pp.508-527.
3. Coyle J.Y., Molliver M.E. and Kuhar M.J., In the situ injection of kainic acid: A new method for selectively lesioning neuronal cell bodies while sparing axons of passage. J. Comp.Neurol. 180(1978)301-324.
4. Gu Y.M., Chen Y.C. and Ye L.M., A new highly sensitive HRP-TMB method using sodium tungstate as a stabilizer: II. Electron microscopic study. Chinese J. Neuroanat. 7(1991)124-129.
5. Lindsley D.F. and Holmes J.E., Basic Human Neurophysiology, Elsevier, New York/Amsterdam/Oxford, 1984, pp62-82.
6. Luo P.F., Wang B.P., Peng Z.Z. and Li J.S., Morphological characteristics and terminating patterns of masseteric neurons of the mesencephalic trigeminal nucleus in the rat: An intracellular horseradish peroxidase labeling study. J. Comp. Neurol.303(1991)286-299.
7. Moriizumi T., Nakamura Y., Okoyama S. and Kitao y., Synaptic organization of the cat entopeduncullar nucleus with special reference to the relationship between the afferents to entopedunculothalamic projection neurons: An electron microscope study by a combined degeneration and horseradish peroxidase tracing technique. Neuroscience 20(1987)797-816.
8. Mugnani E. and Friedrich V.L., Electron microscopy:Identification study of normal and degenerating neuronal elements by electron microscopy. In Heimer L. and Robards M. J. [eds.],Neuroanatomical tract-tracing methods, Plenum Press, New York and London. 1981, pp377-406.
9. Streit W.J. and Kreutzberg G.W., Response of endogenous glial cells to motor neuron degeneration induced by toxic ricin. J.Comp. Neurol. 268(1988)248-263.
10.Takeuchi Y., McLean J.H. and Hopkins D.A., Reciprocal connections between the amygdala and parabrachial nuclei: ultrastructural demonstration by degeneration and axonal transport of horseradish peroxidase in the cat. Brain Research 239(1982)583-588.
11.Wang B.R. and Li J.S., The projection pathway from the trigeminal mesencephalic nucleus (Vme) to the thalamus in the rat. I. The brain stem distribution of the "central" processes of Vme--- A light and electron microscope study. Chinese J.Neuroanat. 4(1988)35-40.
12.Wang B.R. and Li J.S., The projection pathway from the trigeminal mesencephalic nucleus (Vme) to the thalamus in the rat. II. The efferent projection of the dorsal-medial portion of the subnucleus oralis of the trigeminal spinal nucleus(Vodm) and the adjacent lateral reticular formation (LRF). Chinese J.Neuroanat. 4(1988)41-51.
13.Wang B.R. and Li J.S. , The projection pathway from the trigeminal mesencephalic nucleus (Vme) to the thalamus in the rat. III. The projection from the zone-shaped area along the medial border of principal trigeminal nucleus to the thalamus. Chinese J. Neuroanat . 4(1988)53-60.
14.Wiley R.G., Blessing W.W. and Reis D.J., Suicide transport:Destruction of neurons by retrograde transport of ricin, abrin,and modecein. Science 216(1982)889-890.
15.Xie W., Zhang J.D., Yang S. and Li J.S., Relationship between degeneration of mesencephalic trigeminal neurons induced by ricin and different amount of ricin used or different animal survival times. submitted
16.Yamamoto T., Iwasaki Y., Konno H. and kudo H., Retrograde axoplasmic transport of toxic lectins is useful for transganglionic tracings of the peripheral nerve. Brain Research 274(1983)325-328.
17.Zhang J.H., Li H.M. and Li J.S., The relationship between sensitivity, preservation of tissue ultrastructure and the pH of acetate buffer in HRP-TMB reaction medium. Chinese J. Neuroanat.3(1987)118-122.
18.Zhang J.D. and Li J.S., A possible trigeminal proprioceptive pathway from the masseter muscle to the contralateral thalamus---Comprehensive tracing studies in the rat. submitted
19.Zhang J.D. and Li J.S., Nucleus morphology, location,cytoarchitecture and connections of the secondary neurons in the trigeminal proprioceptive pathway in the rat---A light microscopic observation. Chinese J. Neuroanat. 8(1992)63-72.
20.Zhang J.D., Wang B.R., Li H.M. and Li J.S., Projections from neurons innervating the masseter muscle to the subnucleus oralis of the spinal trigeminal nucleus and adjacent lateral reticular formation in the rat. J. Hirnforsch. 32(1991)641-646.
[1] Abe K. A study of sensory projection from jaw muscles to the cerebralcortex inthe rat. Jap J Physiol. 1978;28.309～322
[2] 王百忍，李继硕．大鼠三叉神经中脑核神经元中枢突在脑干内投射的光、电镜观察(中脑核——丘脑投射通路研究之一)．神经解剖学杂志，1988；4：35～40
[3] 王百忍，李继硕．大鼠三叉神经脊束核吻侧亚核背内侧部及邻接的外侧网状结构的传出投射——WGA—HRP顺行追踪研究(中脑核——丘脑投射通路研究之二)．神经解剖学杂志，1988;4，41～52
[4] 王百忍，李继硕．大鼠沿三叉神经感觉主核内缘的带状区向丘脑的投射——HRP法研究(中脑核——丘脑投射通路研究之三)．神经解剖学杂志，1988；4：53～60
[5] Appenteng BK, Donga R, Willians RG. Morphological and electrophysiologicaldetermination of the projections of jaw--elevatormuscle spindle afferents in rats. J Physiol, 1985;369:93～113
[6] Matesz C. Peripheral and cental distribution of fibres of the mesencephalic trigeminal root in the rat. Neurosci Lett, 1981;27:13～17
[7] 罗丕福，李继硕．GABA样神经元和GABA能三叉一丘脑投射神经元在三叉神经感觉核簇的分布及形态学特征．神经解剖学杂志，1991;7：185～194
[8] Rustioni A. Non --primary afferents to the nucleus gracilis fromlumbar cord of the cat. Brain Res, 1973;51:81～95
[9] 吕国蔚．背索突触后神经元．生理科学进展，1984;15：310～315
[10] 罗丕福，李继硕．大鼠三叉神经本体觉三级传导通路的形态学证据——Ricin跨节溃变与HRP逆行追踪结合的光、电镜研究．神经解剖学杂志，1992;8：57～62
[11] Luo PF, Wang BR, Peng ZZ, et al. Morphological characteristics and terminating patterns of masseteric neurons of the mesencephalic trigeminal nucleus in the rat: An intracellular horseradish peroxidase labeling study, j Comp Neurol, 1990(303:286～299
[12] 张敬东，李继硕．大鼠三叉神经本体觉中枢通路二级神经元中继咬肌本体觉传入的超微结构证据．神经解剖学杂志，1992；8：203～210
[13] 李金莲，张敬东，李继硕．三叉神经本体觉中枢通路二、三级神经元之间突触连接的发现及突触构筑学研究——溃变与HRP逆行追踪法相结合的电镜观察．神经解剖学杂志，1993；9：97～104
1．王百忍 李继硕；神经解剖学杂志 4：35，1988
2．王百忍 李继硕：神经解剖学杂志 4：41，1988
3．王百忍 李继硕：神经解剖学杂志 4：53，1988
4．李惠民 李继硕：解剖学报 15：366，1984
5．罗丕福 李继硕：神经解剖学杂志 8：57，1992
6．彭兆知 张凤鸣：解剖学杂志 19(增)：279，1986
7．彭兆知等：神经解剖学杂志 2：27，1986
8. Alvarado-Mallart RM et al, Exp Brain Res 23:167, 1975
9. Appenteng K et al, J Physiol 369:93, 1985
10. Capra NF and Wax TD, J Comp Neurol 279:341, 1989
11. Carpenter MB and Sutin J. Human Neuroanatomy Williams & Wilkins London p 393, 1983
12. Darian-Smith I. Handbook of sensory physiology Springer-Verlag Berlin Heidelb erg New York p27l, 1973
13. Denng-Brian D and Yanagisawa N, Brain 96:7.83, 1973
14. Erzurumlu R and Killackey HP. J Comp Neurol 188:75, 1979
15. Falls WM: Brain Res 290:136, 1981
16. Falls WM, Neuroscience 13:1279, 1984
17. Falls WM et al. Somatosens Res 3:89, 1985
18. Gobel S and Purvis MB. Brain. Res 48:27, 1972
19. Huerta MF et al: J Comp Neurol 220:147, 1983
20. Kelly JP.. Principles of neural science Elsevier New York p562, 1985
21. Kruger L et al: Pain in the trigeminal rigion Elsevier New York pl91, 1977
22. Kruger L et al.. The cranial nerves Springer-Verlag Berlin p273, 1981
23. Luo PF et al, J Comp Neurol 303:286, 1991
24. Matesz C: Neurosci Lett 27:13, 1981
25. Mesulam M-M: Tracing neural connections with horseradish peroxidase John Wiley & Sons New York p 1 , 1982
26. Mizuno N et al. J Comp Neurol 215:290, 1983
27. Nishimori T et al. Brain Res 372:375, 1986
28. Nomura S and Mizuno N. Neurosci Lett Suppl 17:sl52, 198.)
29. Ruggireo DA et al: Brain Res 225:225, 1981
30. Schvarcz JR: Advances in pain research and therapy Raven. Press New York p 331, 1979
31. Shigenaga Y et al: J Comp Neurol 243:388, 1986
32. Shigenaga Y et al. J Comp Neurol 244:1, 1986
33. Shigenaga Y et al: J Comp Neurol 268:489, 1988
34. Zhang JD et al. J Hirnforsch 32:641, 1991
1．王百忍，李继硕．大鼠三叉神经中脑核神经元中枢突在脑干内投射的光、电镜观察(中脑核——丘脑投射通路研究之一)。神经解剖学杂志，1988；4：35-40
2．王百忍，李继硕．大鼠三叉神经脊束核吻侧亚核背内侧部及邻近的外侧网状结构的传出投射——WGA—HRP顺行追踪研究(中脑核——丘脑投射通路研究之二)。神经解剖学杂志，1988；4：41-52
3．王百忍，李继硕．大鼠沿三叉神经感觉主核内缘的带状区向丘脑的投射——HRP法研究(中脑核——丘脑投射通路研究之三)，神经解剖学杂志，1988；4：53～60
4．顾耀铭，陈以慈，叶鹿鸣．钨酸钠作为稳定剂的新的高更敏HRP-TMB法——Ⅱ。电镜研究。神经解剖学杂志，1991；7：124-129
5. Mesulum MM. Tetramethyl benzidine for horseradish peroxidase neuro-histochemistry. A non-carcinogenic blue reaction product with superior sensitivity for visualizing neural afferents and efferents. J Histochem Cytochem, 1978; 26:106-117
6．张敬东，李继硕．三叉神经本体觉中枢通路二级神经元所在地(三叉神经脊束核吻侧亚核背内侧区，Vodm)的核团形态、细胞构筑及纤维联系——光镜观察．神经解剖学杂志，1992；8：633-72
7. Faxinos G, Watson C. The rat. brain in stereotaxic coordinates, 2nd ed. Academic Press, Sydney/Orlando/San Diego/NewYork/Austin/London/Montreal/Tokyo/Toronto, 1986.
8．王振福，………李继硕．大鼠沿三叉神经感觉主核内缘的“带状区”向丘脑的投射和细胞构筑-HRP法研究．神经解剖学杂志，1995(在印刷中)
9. Carpenter MB, Sutin J. Human Neuroanatomy, 8th ed. Baltimore/London, Williams & Wilkins, 1983, pp.516-524
10. Beckstead RM, Morse JR, Norgren R. The nucleus of the solitary tractin the monkey:Projections to the thalamus and brain stem nuclei. J Comp Neurol, 1980; 190:259-282
11. Faull RLM, Mehler WR. Thalamus. In G Paxinos (ed.), The rat nervous system. Vol 1,Academic Press, Sydney, 1985, pp. 129-168
12. Smith RL. The ascending fiber projections from the principal sensory trigeminal nucleus in the rat. J Comp Neurol, 1973; 148:423-446
13. Smith RL. Axonal projections and connections of the principal sensory trigeminal nucleus in the monkey. J Comp Neurol, 1975; 163:347-376
14. Fukushima I, Kerr FWL. Organization of trigeminothalamic tracts and other thalamic afferent systems of the brainstem in the rat: Presence of gelatinosa neurons with thatamic connections. J Comp Neurol, 1979; 183:169-184
15. Eisenman J, Fromm G, Landgren S, et al. The ascending projections of trigeminal neurones in the cat, investigated by antidromic stimulation. Acta Physiol Scand, 1964; 60:337-350
16. Jerge CR. The organization and function of the trigeminal mesencephalic nucleus. J Neurophysiol, 1963; 26:379-392
17. Lund JP, Sessle BJ. Oral-facial and jaw muscle afferent projections to neurons in cat frontal cortex. Exp Neurol, 1974; 45:314-331
18. Abe K. A study of sensory projection from jaw muscles to the cerebral cortex in the rat. Jap J Physiol, 1978; 28:309-322
[1] Matesz C. Peripheral and central distribution of fibres of the mescnccphalic trigcminal root in the rat. Ncurosci Lett, 1981;27:13～17
[2] 彭兆知，曹广义，李继硕．大鼠三叉神经中脑核神经元中枢突的行径和分布——HRP跨节追踪研究．神经解剖学杂志，1986;2：27～36
[3] Juch PJW, Roxk JTM. Efferent projections of the parvocellular reticular nucleus to the mesencephalic trigeminal nucleus in rat. Brain Res, 1988;462:185～189
[4] Roxk JTM, Juch PJW, van Willigen JD. Arrangement and connections of mesencephalic trigeminal neurons in the rat. Acta Anat, 1986;127:7～15
[5] Zhang JD, Wang BR, Li HM, et al. Projections from neurons innervating the masscter muscle to the subnuclcus oralis of the spinal trigeminal nucleus and adjacent lateral reticular formation in the rat. J Hirnforsch, 1991;32:641~646
[6] 王百忍，李继硕．大鼠三叉神经中脑核神经元中枢突在脑干内投射的光、电镜观察(中脑核——丘脑投射通路研究之一)．神经解剖学杂志，1988;4：35～40
[7] 王百忍，李继硕．大鼠三叉神经脊束核吻侧亚核背内侧部及邻接的外侧网状结构的传出投射——WGA-HRP顺行追踪研究(中脑核——丘脑投射通路研究之二)．神经解剖学杂志，1988；4：41～52
[8] 王百忍，李继硕．大鼠沿三叉神经感觉主核内缘的带状区向丘脑的投射——HRP法研究(中脑核——丘脑投射通路研究之三)．神经解剖学杂志，1988；4：53～60
[9] 张敬东，李继硕．大鼠三叉神经本体觉中枢通路二级神经元中继咬肌本体觉传入的超微结构证据．神经解剖学杂志，1992；8：203～210
[10] 李金莲，张敬东，李继硕．三叉神经本体觉中枢通路二、三级神经元之间突触连接的发现及突触构筑学研究——溃变与HRP逆行追踪法相结合的电镜观察．神经解剖学杂志，1993;9：97～104
[11] 张敬东，李继硕．三叉神经本体觉中枢通路二级神经元所在地(三叉神经脊束核吻侧亚核背内侧区，Vodm)的核团形态、细胞构筑及纤维联系——光镜观察神经解剖学杂志，1992;8：63～72
[12] 顾耀铭，陈以慈．叶鹿鸣．钨酸钠作为稳定剂的新的高灵敏HRP-TMB法——Ⅱ、电镜研究．神经解剖学杂志，1991；7：124～129
[13] Roxk JTM, Luiten PGM, van Willigen JD. Afferent projections to the mesencephalic trigeminal nucleus in the rat. Acta Anat, 1988;132:269～274
[14] Hinrichsen CFL, Larramendi LMH?Synapses and cluster formation in the mouse mesencephalic fifth nucleus. Brain Res,1968;7:296～299
[15] Hinrichsen CFL, Larramendi LMH. The trigeminal mesencephalic nucleus IⅠ. Electron microscopy. Am J Anat, 1970;127:303～320
[16] Liem RSB, Copray JCVM , van Willigen JD. Ultrastructure of the rat mesencephalic trigeminal nucleus. Acta Anat, 1991;140:112～119
[17] Inagaki N, Yamatodani A, Shinoda K, et al. The histaminergic innervation of mesencephalic neurons of the trigeminal nerve in rat brain: a light and electron microscopical study. Brain Res, 1987;418:388～391
[18] Copray JCVM, Liem RSB, Ter Horst GJ, et al. Dopaminergic afferents to the mesencephalic trigeminal nucleus: a light and electron microscope immunocytochemistry study. Brain Res, 1990;541:343～348
[19] Copray JCVM, Ter Horst GJ, Liem RSB, et al. Neurotransmitters and neuropeptides within the mesencephalic trigeminal nucleus of the rat: An immunohistochemical analysis. Neuroscience, 1990;37:399～411
[20] 史中立，秦秉志，李继硕．大鼠骶髓后连合核的中枢联系及机能分析——HRP法研究．解剖学报，1990;21：360～368
[21] 贾宏阁，饶志仁，施际武．大鼠臂旁核内接受孤束核内脏传入之神经元同时接受中央杏仁核的反馈调节——溃变、HRP法结合包埋后免疫电镜研究．神经解剖学杂志，1994；10：294～298
[22] Kayahara T. Synaptic connections between spinal motoneurons and dorsal root ganglion cells in the cat. Brain Res, 1986; 376:299～309
[1] Hinrichsen CFL, Larrnmendi LMH. Synapses and cluster formation in the mouse metiencephalic fifth nucleus. Brain Res,1968;7:296～299
[2] Hinrichsen CFL, Laxramendi LMH. The trigeminal mesencephalic nucleus IⅠ . Electron microscopy, Arm J Anat,1970;127:303～320
[3] Liem RSB, Copray JCVM, van Willigen JD. Ultrastructure of the rat mesencephalic trigerminal nucleus. Acta Anat,1991;140:112～119
[4] Inagaki N, Yamatodani A, Shinoda K, et al. The histaminergic innervation of mesencephalic neurons of the trigeminal nerve in rat brain : a light and electron microscopical study. Brain Res, 1987 ; 418:388～391
[5] Copray JCVM , Liem RSB. Ter Horst GJ, et al. Dopaminergic afferents to the mesencephalic trigeminal nucleus; a light and electron microscope immunocytochemistry study. Brain Res. 1990 ; 514 : 343～348
[6] Copray JCVM. Ter Horst GJ, Liem RSB. et al. Neurotransmitters and neuropeptides within the mesencephalic trigeminal nucleus of the rat; an immunohistochemical analysis. Neuroscience. 1990;37: 399～411
[7] Carpenter MB, Sutin J. Hur an neuroanatomy. Baltimore'London : Williams & Wilkins, 1983:393～401
[8] Kayahara T, Takimoto T, Sakashita S. Synaptic junctions in the cat spinal ganglion. Brain Res, 1981; 216:277～290
[9] Kayahara T, Sakashita S, Takimoto T. Evidence for spinal origin of neurons synapsing with dorsal root ganglion cells of the cat. Brain Res. 1984;293:225～230
[10] Kayahara T, Synaptic connections between spinal motonenrons and dorsal root ganglion cells in the cat. Brain Res, 1986;376:299～309
[11] 李云庆，饶志仁，施际武．大鼠三叉神经感觉核簇内SP受体及三叉神经节内SP样阳性终末和SP受体分布的免疫组织化学研究．神经解剖学杂志，1993；9：89～92
[12] Peters A. Palay SL, de E Webster H. The fine structure of the nervous system. London: WB Saunders Company. 1976;118～180
[13] De Montigny C, Lund JP. A microiontophoretic study of the action of kainic acid and putative neurotransmitters in the rat Tnesenvephalic trigeminal nucleus. Neuroscience , 1980;5 ,1621～1628
[14] Guyenet PG, Aghajanian GK. Excitation of neurons in the locus coeruleus by substance P and related peptide. Brain Res, 1977;136:178～184
[15] Henderson G, Pepper CM, Shefner SA. Electrophysiological properties of neurons contained in the locus coeruleus and mesencephalic nucleus of the trigeminal nerve in vitro. Exp Brain Res, 1982;45;29～37
[16] Minkels RF, Juch PJW, Ter Horst GJ, et al. Projections of the parvocellular reticular formation to the contralaterai mesencephalic trigeminal nucleus in the rat. Brain Res, 1991;547:13～21
[17] Liem RSB, Copray JCVM, van Willigen JD. Distribution of synaptic boutons in the mesencephalic trigeminal nucleus of the rat -- A quantitative electron-microscopical study. Acta Anat, 1992; 143; 74～78
[18] Luo PF. Wang BR, Peng ZZ. et al. Morphological characteristics and terminating patterns of masseteric neurons of the mesencephalic trigeminal nucleus in the rat: An intracellular horseradish peroxidase labeling study. J Comp Neurol, 1991 ; 303:286～299
[19] Shigenaga Y. Mitsuhiro Y, Yoshida A. et al. Morphology of single mesencephalic trigeminial neurons innervating masseter muscle of the cat. Brain Res, 1988;445:392～399
[20] Shigenaga Y, Mitsuhiro Y, Shirana Y , et al. Two types of jaw-muscle spindle afferents in the cat as demonstrated by intraaxonal staining with HRP. Brain Res. 1990;514: 219～237