Involvement of alpha-2 adrenoceptors in the periaqueductal gray-induced inhibition of dorsal horn cell activity in rats

Journal of Pharmacology and Experimental Therapeutics

Volumn 278, Issue 1, 1996, Pages 125-135

  Peng, Yuan B ^a   Lin, Qing ^a   Willis, William D ^a  

^a UNIVERSITY OF TEXAS MEDICAL BRANCH   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

ALPHA 2 ADRENERGIC RECEPTOR; CLONIDINE; IDAZOXAN; NORADRENALIN; YOHIMBINE;

ANALGESIA; ANIMAL EXPERIMENT; ANTINOCICEPTION; ARTICLE; CONTROLLED STUDY; DOSE RESPONSE; ELECTROSTIMULATION; LOCUS CERULEUS; MECHANICAL STIMULATION; MICRODIALYSIS; NONHUMAN; NORADRENERGIC SYSTEM; PARABRACHIAL NUCLEUS; PERIAQUEDUCTAL GRAY MATTER; PRIORITY JOURNAL; RAT; SINGLE UNIT ACTIVITY; SPINAL CORD DORSAL HORN;

ANIMALS; CLONIDINE; DOSE-RESPONSE RELATIONSHIP, DRUG; GANGLIA, SPINAL; PERIAQUEDUCTAL GRAY; RATS; RATS, SPRAGUE-DAWLEY; RECEPTORS, ADRENERGIC, ALPHA-2; SPINAL CORD; YOHIMBINE;

EID: 0030440934     PISSN: 00223565     EISSN: None     Source Type: Journal    
DOI: None     Document Type: Article

References (144)

1. AIMONE, L. D. AND GEBHART, G. F.: Spinal monoamine mediation of stimulation-produced antinociception from the lateral hypothalamus. Brain Res. 403: 290-300, 1987.
2. AIMONE, L. D., JONES, S. L. AND GEBHART, G. F.: Stimulation-produced descending inhibition from the periaqueductal gray and nucleus raphe magnuis in the rat: Mediation by spinal monoamines but not opioids. Pain 31: 123-136, 1987.
3. ARCHER, T., ARWESTREM, E., JONSSON, G., MINOR, G. AND POST, C.: Noradrenergicserotonergic interactions and nociception in the rat. Eur. J. Pharmacol. 120: 295-307, 1986.
4. ASTON-JONES, G., ENNIS, M., PIERIBONE, V. A., NICKELL, W. T. AND SHIPLEY, M. T.: The brain nucleus locus coeruleus: Restricted afferent control of a broad efferent network. Science (Wash. DC) 234: 734-737, 1986.
5. ASTON-JONES, G., SHIPLEY, M. T., CHOUVET, G., ENNIS, M., VAN BOCKSTAELE, E., PIERIBONE, V., SHIEKHATTAR, R., AKAOKA, H., DROLET, G., ASTIER, B., CHARLETY, P., VALENTINO, R. J. AND WILLJAMS, J, T.: Afferent regulation of locus coeruleus neurons: Anatomy, physiology and pharmacology. Prog. Brain Res. 88: 47-75, 1991.
6. BASBAUM, A. I., CLANTON, C. H. AND FIELDS, H. L.: Three bulbospinal pathways from the rostral medulla of the cat: An autoradiographic study of pain modulating systems. J. Comp. Neurol. 178: 209-224, 1978.
7. BASBAUM, A. I. AND FIELDS, H. L.: Endogenous pain control mechanisms: Review and hypothesis. Ann. Neurol. 4: 451-462, 1978.
8. BASBAUM, A. I. AND FIELDS, H. L.: The origin of descending pathways in the dorsolateral funiculus of the spinal cord of the cat and rat: Further studies on the anatomy of pain modulation. J. Comp. Neurol. 187: 513-532, 1979.
9. BASBAUM, A. I. AND FIELDS, H. L.: Endogenous pain control systems: Brainstem spinal pathways and endorphin circuitry. Ann. Rev. Neurosci. 7: 309-338, 1984.
10. BASBAUM, A. I., GLAZER, E. J. AND OERTEL, W.: Immunoreactive glutamic acid decarboxylase in the trigeminal nucleus caudalis of the cat: A light- and electron-microscopic analysis. Somatosens. Motor Res. 4: 77-94, 1986.
11. BEALL, J. E., MARTIN, R. F., APPLEBAUM, A. E. AND WILLIS, W. D.: Inhibition of primate spinothalamic tract neurons by stimulation in the region of the nucleus raphe magnus. Brain Res. 114: 328-333, 1976.
12. BEITZ, A., MULLETT, M. A. AND BRANDT, N.: The relationship of periaqueductal gray projections to bulbospinal neurons: A combined fluorogold-PHA-L analysis. Soc. Neurosci. Abstr. 14: 856, 1988.
13. BBITZ, A. J.: The organization of afferent projections to the midbrain periaqueductal gray of the rat. Neuroscience 7: 133-159, 1982.
14. BELCHER, G., RYALL, R. W. AND SCHNAFFNER, R.: The differential effects of 5-HT, noradrenaline and raphe stimulation on nociceptive and non-nociceptive dorsal horn interneurons in the cat. Brain Res. 151: 307-321, 1978.
15. BERVOETS, K. AND MILLAN, M. J.: 5-HT1A receptors and the tail-flick response. V. Opposite modulation of 5-HT1A receptor-induced spontaneous tail-flicks by alpka-1A as compared with alpka-2D adrenoceptors in rat lumbar spinal cord. J. Pharmacol. Exp. Ther. 269:110-120, 1994.
16. BESSON, J. M., GUILBAUD, G. AND LEBARS, D.: Descending inhibitory influences exerted by the brainstem upon the activities of dorsal horn lamina V cells induced by intra-arterial bradykinin into the limbs. J. Physiol. (Lond.) 248: 725-739, 1975.
17. BOBILLIEE, P., SEGUIN, S., PETITJEAN, F., SALVERT, D., TOURET, M. AND JOUVET, M.: The raphe nuclei of the cat brain stem: A topographical atlas of their efferent projections as revealed by autoradiography. Brain Res. 113: 449-486, 1976.
18. BONNET, F., BOICO, O., ROSTAING, S., LOREFERNE, J. F. AND SAADA, M.: Clonidineinduced analgesia in postoperative patients: Epidural versus intramuscular administration. Anesthesiology 72: 423-427, 1990.
19. 3H]-idazoxan in the human cerebral cortex. Neurochem. Int. 22: 153-163, 1993.
20. BROWN, A. G.: Effects of descending impulses on transmission through the spinocervical tract. J. Physiol. (Lond.) 219: 103-125, 1971.
21. BYRUM, C. E. AND GUYENET, P. G.: Afferent and efferent connections of the A5 noradrenergic cell group in the rat. J. Comp. Neurol. 261: 529-542, 1987.
22. CAMERON, A. A., KHAN, I. A., WESTLUND, K. N. AND WILLIS, W. D.: The efferent projections of the periaqueductal gray in the rat: A Phaseoleus vulgarisleucoagglutinin study. II. Descending projections. J. Comp. Neurol. 351: 585-601, 1995.
23. CARLTON, S. M., HONDA, C. N., WILLCOCKSON, W. S., LACRAMPE, M., ZHANG, D., DENOROY, L., CHUNG, J. M. AND WILLIS, W. D.: Descending adrenergic input to the primate spinal cord and its possible role in modulation of spinothalamic cells. Brain Res. 543: 77-90, 1991.
24. CEDARBAUM, J. M. AND AGHAJANIAN, G. K.: Afferent projections to the rat locus coeruleus as determined by a rertrograde tracing technique. J. Comp. Neurol. 178: 1-16, 1978.
25. CERVERO, F., IGGO, A. AND OGAWA, H.: Nociceptor-driven dorsal horn neurones in the lumbar spinal cord of the cat. Pain 2: 5-24, 1976.
26. CHUNG, J. M., SURMEIER, D. J., LEE, K. H., SORKIN, L. S., HONDA, C. N., TSONG, Y. AND WILLIS, W. D.: Classification of primate spinothalamic and somatosensory thalamic neurons based on cluster analysis. J. Neurophysiol. 56: 308-327, 1986.
27. CHUNG, R. Y., MASON, P., STRASSMAN, A. AND MACIEWICZ, R.: Suppression of the jaw-opening reflex by periaqueductal gray stimulation is decreased by paramedian brainstem lesions. Brain Res. 403: 172-176, 1987.
28. CLARK, F. M. AND PROUDFIT, H. K.: The projection of locus coeruleus neurons to the spinal cord in the rat determined by anterograde tracing combined with immunocytochemistry. Brain Res. 538: 231-245, 1991a.
29. CLARK, F. M. AND PROUDFIT, H. K.: Projections of neurons in the ventromedial medulla to pontine catecholamine cell groups involved in the modulation of nociception. Brain Res. 540: 105-115, 1991b.
30. CLARK, F. M. AND PROUDFIT, H. K.: Anatomical evidence for genetic differences in the innervation of the rat spinal cord by noradrenergic locus coeruleus neurons. Brain Res. 591: 44-53, 1992.
31. CLARK, F. M. AND PROUDFIT, H. K.: The projections of noradrenergic neurons in the A5 catecholamine cell group to the spinal cord in the rat: Anatomical evidence that A5 neurons modulate nociception. Brain Res. 616: 200-210, 1993.
32. CURTIS, D. R., LEAH, J. D. AND PEET, M. J.: Spinal interneuron depression by DS103-282. Br. J. Pharmacol. 79: 9-11, 1983.
33. DAHLSTRÖM, A. AND FUXE, K.: Evidence for the existence of monoamine neurons in the central nervous system: The distribution of monoamine terminals in the central nervous system. Acta Physiol. Scand. 247: suppl. 64, 37-84, 1965.
34. DANZEBRINK, R. M. AND GEBHART, G. F.: Intrathecal coadministration of clonidine with serotonin receptor agonists produces supra-additive visceral antinociception in the rat. Brain Res. 555: 35-42, 1991.
35. DAVIES, J.: Effects of tizanidine, eperisone and afloqualone on feline dorsal horn neuronal responses to peripheral cutaneous noxious and innocuous stimuli. Neuropharmacology 28: 1357-1362, 1989.
36. DAVIES, J. AND QUINLAN, J. E.: Selective inhibition of responses in feline dorsal horn neurons to noxious cutaneous stimuli by tizanidine (DS 103-282) and noradrenaline. Involvement of alpha-adrenoceptors. Neuroscience 16: 673-682, 1985.
37. DICKHAUS, H., PAUSER, G. AND ZIMMERMANN, M.: Tonic descending inhibition affects intensity coding of nociceptive responses of spinal dorsal horn neurones in the rat. Pain 23:145-158, 1985.
38. DOUGHERTY, P. M., PALECEK, J. AND PALECKOVA, V.: Infusion of substance P or neurokinin A by microdialysis alters responses of primate spinothalamic tract neurons to cutaneous stimuli and to iontophoretically released excitatory amino acids. Pain 61: 411-425, 1995.
39. DRASNER, K. AND FIELDS, H. L.: Synergy between the antinociceptive effects of intrathecal clonidine and systemic morphine in the rat. Pain 32: 309-312, 1988.
40. DUGGAN, A. W., GRIERSMITH, B. T. AND JOHNSON, S. M.: Supraspinal inhibition of the excitation of dorsal horn neurones by impulse in unmyelinated primary afferents: Lack of effect by strychnine and bicuculline. Brain Res. 210: 231-241, 1981.
41. DUGGAN, A. W., HALL, J. G., HEADLEY, P. M. AND GRIERSMITH, B. T.: The effect of naloxone on the excitation of dorsal horn neurons of the cat by noxious and non-noxious cutaneous stimuli. Brain Res. 138: 185-189, 1977.
42. ENNIS, M. AND ASTON-JONES, G.: GABA-mediated inhibition of locus coeruleus from the dorsomedial rostral medulla. J. Neurosci. 9: 2973-2981, 1989.
43. ENNIS, M., BEHBEHANI, M., SHIPLEY, M. T., VAN BOCKSTAELE, E. J. AND ASTON-JONES, G.: Projections from the periaqueductal gray to the rostromedial pericoerulear region and nucleus locus coeruleus: Anatomic and physiologic studies. J. Comp. Neurol. 306: 480-494, 1991.
44. 2-adrenoceptors in the ventrolateral medulla. Eur. J. Pharmacol. 134: 1-13, 1987.
45. FLEETWOOD-WALKER, S. M., MITCHELL, R., HOPE, P. J., MOLONY, V. AND IGGO, A.: An alpha-2 receptor mediates the selective inhibition by noradrenaline of nociceptive responses of identified dorsal horn neurons. Brain Res. 334: 243-254, 1985.
46. FRITSCKY, J. M. AND GRZANNA, R.: Demonstration of two separate descending noradrenergic pathways to the rat spinal cord: Evidence for an intragriseal trajectory of locus coeruleus axons in the superficial layers of the dorsal horn. J. Comp. Neurol. 291: 553-582, 1990.
47. FRITSCHY, J. M. AND GRZANNA, R.: Restoration of ascending noradrenergic projections by residual locus coeruleus neurons: Compensatory response to nerotoxin-induced cell death in the adult rat brain. J. Comp. Neurol. 321: 421-441, 1992.
48. FRITSCHY, J. M., LYONS, W. E., MULLEN, C. A., KOSOFSKY, B. E., MOLLIVER, M. E. AND GEZANNA, R.: Distribution of locus coeruleus axons in the rat spinal cord: A combined anterograde transport and immunohistochemical study. Brain Res. 437: 176-180, 1987.
49. FU, L., SHIPLEY, M. T. AND ASTON-JONES, G.: Dendrites of rat locus coeruleus neurons are asymmetrically distributed: Immunocytochemical LM and EM studies. Soc. Neurosci. Abstr. 15: 1013, 1989.
50. Fu, L., SHIPLEY, M. T., ASTON-JONES, G., CHIANG, C. AND PIERIBONE, V. A.: Architecture of the pericoerulear region in the rat. Soc. Neurosci. Abstr. 14: 404, 1988.
51. GEBHART, G. F.: Opiate and opioid peptide effects on brainstem neurons: Relevance to nociception and antinociceptive mechanisms. Pain 12: 93-140, 1982.
52. GEBHART, G. F.: Modulatory effects of descending systems on spinal dorsal horn neurons. In Spinal Afferent Processing, ed. by T. L. Yaksh, pp. 391-416, Plenum Press, New York, 1986.
53. GEBHAHT, G. F., SANDKÜHLER, J., THALHAMMER, J. G. AND ZIMMERMAN, M.: Inhibition of spinal nociceptive information by stimulation in midbrain of the cat is blocked by lidocaine microinjected in nucleus raphe magnus and medullary reticular formation. J. Neurophysiol. 50: 1446-1458, 1983.
54. GEBHART, G. F., SANDKÜHLER, J., THALHAMMER, J. G. AND ZIMMERMANN, M.: Inhibition in spinal cord of nociceptive information by electrical stimulation and morphine microinjection at identical sites in midbrain of the cat. J. Neurophysiol. 51: 75-89, 1984.
55. GERHART, K. D., YEZIERSKI, R. P., WILCOX, T. K. AND WILLIS, W. D.: Inhibition of primate spinothalamic tract neurons by stimulation in periaqueductal gray or adjacent midbrain reticular formation. J. Neurophysiol. 51: 450-466, 1984.
56. 3H]serotonin are presynaptic to enkephalin immunoreactive neurons in cat dorsal horn. Brain Res. 298: 386-391, 1984.
57. GLAZER, E. J. AND BASBAUM, A. I.: Immunohistochemical localization of leucineenkephalin in the spinal cord of the cat: Enkephalin-containing marginal neurons and pain modulation. J. Comp. Neurol. 196: 377-389, 1981.
58. GORDH, T.: Epidural clonidine for treatment of postoperative pain after thoracotomy: A double-blind placebo-controlled study. Acta Anaesthesiol. Scand. 32: 702-709, 1988.
59. GRZANNA, R. AND FRITSCHY, J. M.: Efferent projections of different subpopulations of central noradrenaline neurone. Prog. Brain Res. 88: 89-101, 1991.
60. HAMMOND, D. L.: Control systems for nociceptive afferent processing: The descending inhibitory pathways. In Spinal Afferent Processing, ed. by T. L. Yaksh, pp. 363-390, Plenum, New York, 1986.
61. HAMMOND, D. L., TYCE, G. M. AND YAKSH, T. L.: Efflux of 5-hydroxytryptamine and noradrenaline into spinal cord superfusates during stimulation of the rat medulla. J. Physiol. (Lond.) 359: 151-162, 1985.
62. HANDWERKER, H. O., IGGO, A. AND ZIMMERMANN, M.: Segmental and supraspinal actions on dorsal horn neurons responding to noxious and non-noxious skin stimuli. Pain 1:147-165, 1975.
63. HAYES, R. L., PRICE, D. D., RUDA, M. AND DUBNER, R.: Suppression of nociceptive responses in the primate by electrical stimulation of the brain or morphine administration: Behavioral and electrophysiological comparisons. Brain Res. 167:417-421, 1979.
64. HEADLEY, P. M., DUGGAN, A. W. AND GRIERSMITH, B. T.: Selective reduction of noradrenaline and 5-hydroxytryptamine of nociceptive responses of cat dorsal horn neurones. Brain Res. 145: 185-189, 1978.
65. HEINRICHER, M. M., HAWS, C. M. AND FIELDS, H. L.: Opposing actions of norepinephrine and clonidine on single pain-modulating neurons in rostral ventromedial medulla. In Proceedings of the 5th World Congress on Pain, pp. 590-694, Elsevier, Amsterdam, 1988.
66. HILLMAN, P. AND WALL, P. D.: Inhibitory and excitatory factors influencing the receptive fields of lamina 5 spinal cord cells. Exp. Brain Res. 9: 284-306, 1969.
67. HOLSTEGE, G. AND KUYPERS, H. G. J. M.: The anatomy of brainstem pathways to the spinal cord in cat: A labeled amino acid tracing study. Prog. Brain Res. 57: 45-76, 1982.
68. HOWE, J. R. AND ZIEGLGÄNSBERGER, W.: Responses of rat dorsal horn neurons to natural stimulation and to iontophoretically applied norepinephrine. J. Comp. Neurol. 255: 1-17, 1987.
69. HYLDEN, J. L. K., THOMAS, D. A., IADAROLA, M. J., NAHIN, R. L. AND DUBNER, R.: Spinal opioid analgesic effects are enhanced in a model of unilateral inflammation/hyperalgesia: Possible involvement of noradrenergic mechanisms. Eur. J. Pharmacol. 194: 135-143, 1991.
70. KWIAT, G. AND BASBAUM, A. I.: Organization and cytochemistry of brainstem neurons with divergent collaterals to the midbrain periaqueductal gray and the spinal cord. Soc. Neurosci. Abstr. 13: 302, 1987.
71. 2-adrenergic binding sites. Mol. Pharmacol. 37: 876-885, 1990.
72. LEVITT, P. AND MOORE, R. Y.: Origin and organization of brainstem catecholamine innervation in the rat. J. Comp. Neurol. 186: 505-528, 1979.
73. A antagonists reduce the inhibition of primate spinothalamic tract neurons produced by stimulation in periaqueductal gray. Brain Res. 654: 286-302, 1994.
74. 1A receptors. J. Pharmacol. Exp. Ther. 276: 958-967, 1996.
75. MAGOUL, R., ONTENIENTE, B., GEFFARD, M. AND CALAS, A.: Anatomical distribution and ultrastructural organization of the GABAergic system in the rat spinal cord: An immunocytochemical study using anti-GABA antibodies. Neuroscience 20: 1001-1009, 1987.
76. MANTYH, P. W.: Connections of midbrain periaqueductal gray in the monkey. I. Ascending efferent projections. J. Neurophysiol. 49: 567-581, 1983.
77. MANTYH, P. W. AND PESCHANSKI, M.: Spinal projections from the periaqueductal gray and dorsal raphe in the rat, cat and monkey. Neuroscience 7: 2769-2776, 1982.
78. MARCHAND, J. E. AND HAGINO, N.: Afferents to the periaqueductal gray in the rat: A horseradish peroxidase study. Neuroscience 9: 95-106, 1983.
79. MASON, P. AND FIELDS, H. L.: Axonal trajectories and terminations of on- and off-cells in the lower brainstem of the cat. J. Comp. Neurol. 288: 185-207, 1989.
80. MAX, M. B., SCHAFER, S. C., CULNANE, M., DUBNER, R. AND GRACELY, R. H.: Association of pain relief with drug side effects in postherpetic neuralgia: A single-dose study of clonidine, codeine, ibuprofen and placebo. Clin. Pharmacol. Ther. 43: 363-371, 1988.
81. MAYER, D. J. AND PRICE, D. D.: Central nervous system mechanism of analgesia. Pain 2: 379-404, 1976.
82. MCMAHON, S. B. AND WALL, P. D.: Descending excitation and inhibition of spinal cord lamina I projection neurons. J. Neurophysiol. 59: 1204-1219, 1988.
83. MELLER, S. T. AND DENNIS, B. J.: Afferent projections to the periaqueductal gray in the rabbit. Neuroscience 19: 927-964, 1986.
84. MICHEL, M. C. AND ERNSBERGER, P.: Keeping an eye on the I site: Imidazolinepreferring receptors. Trends Pharmacol. Sci. 13: 369-370, 1992.
85. MITCHELL, K., SPIKE, R. C. AND TODD, A. J.: An immunocytochemical study of glycine receptors and GABA in laminae I-III of rat spinal dorsal horn. J. Neurosci. 13: 2371-2381, 1993.
86. MONROE, P. J., SMITH, D. L., KIRK, H. R. AND SMITH, D. J.: Spinal noradrenergic imidazoline receptors do not mediate the antinociceptive action of intrathecal clonidine in the rat. J. Pharmacol. Exp. Ther. 273: 1057-1062, 1995.
87. NAKAGAWA, I., OMOTE, K., KITAHATA, L. M., COLLINS, J. G. AND MURATA, K.: Serotonergic mediation of spinal analgesia and its interaction with noradrenergic systems. Anesthesiology 73: 474-478, 1990.
88. NESS, T. J. AND GEBHART, G. F.: Quantitative comparison of inhibition of visceral and cutaneous spinal nociceptive transmission from the midbrain and medulla in the rat. J. Neurophysiol. 58: 850-865, 1987.
89. NESS, T. J. AND GEBHART, G. F.: Differential effects of morphine and clonidine on visceral and cutaneous nociceptive transmission in the rat. J. Neurophysiol. 62: 220-230, 1989.
90. NORTH, R. A. AND YOSHIMURA, M.: The actions of noradrenaline on neurones of the rat substantia gelatinose in vitro. J. Physiol. (Lond.) 349: 43-55, 1984.
91. OMOTE, K., KITAHATA, L. M., COLLINS, J. G., NAKATANI, K. AND NAKAGAWA, I.: Interaction between opiate subtype and alpha-2 adrenergic agonists in suppression of noxiously evoked activity of WDR neurons in the spinal dorsal horn. Anesthesiology 74: 737-743, 1991.
92. OSSIPOV, M. H., HARRIS, S., LLOYD, P. AND MESSINEO, E.: An isobolographic analysis of the antinociceptive effect of systemically and intrathecally administered combinations of clonidine and opiates. J. Pharmacol. Exp. Ther. 255: 1107-1116, 1990a.
93. OSSIPOV, M. H., HARRIS, S., LLOYD, P., MESSINEO, E., LIN, B.-S. AND BAGLEY, J.: Antinociceptive interaction between opioids and medetomidine: Systemic additivity and spinal synergy. Anesthesiology 73: 1227-1235, 1990b.
94. PAXINOS, G. AND WATSON, C.: The Rat Brain in Stereotaxic Coordinates, Academic Press, New York, 1986.
95. 2-adrenoceptors in the periaqueductal gray-induced inhibition of dorsal horn cell activity in rats. Soc. Neurosci. Abstr. 21: 1172, 1995.
96. 3 receptors in periaqueductal gray-induced inhibition of nociceptive dorsal horn neurons in rats. J. Pharmacol. Exp. Ther. 276: 116-124, 1996.
97. PRIESTLY, J. V. AND CUELLO, A. C.: Ultrastructural and neurochemical analysis of synaptic input to trigeminothalamic projection neurons in lamina I of the rat: A combined immunocytochemical and retrograde labelling study. J. Comp. Neurol. 285: 467-486, 1989.
98. PRIETO, G. J., CANNON, J. T. AND LIEBESKIND, J. C.: Nucleus raphe magnus lesions disrupt stimulation-produced analgesia from ventral but not dorsal midbrain areas in the rat. Brain Res. 261: 53-57, 1983.
99. PROUDFIT, H. K: Pharmacologic evidence for the modulation of nociception by noradrenergic neurons. Prog. Brain Res. 77: 357-370, 1988.
100. PUKE, M. J. C., Xu, X.-J. AND WIESENFELD-HALLIN, Z.: Intrathecal administration of clonidine suppresses autotomy, a behavioral sign of chronic pain in rats after sciatic nerve section. Neurosci. Lett. 133:199-202, 1991.
101. REYNOLDS, D. V.: Surgery in the electrical analgesia induced by focal brain stimulation. Science (Wash. DC) 164: 444-445, 1969.
102. RUDA, M. A., ALLEN, B. AND GOBEL, S.: Ultrastructural analysis of medial brain stem afferents to the superficial dorsal horn. Brain Res. 205:175-180, 1981.
103. RUDA, M. A., BENNETT, G. J. AND DUBNER, R.: Neurochemistry and neural circuitry in the dorsal horn. Prog. Brain Res. 66: 219-268, 1986.
104. RUDA, M. A., COFFIELD, J. AND DUBNER, R.: Demonstration of postsynaptic opioid modulation of thalamic projection neurons by the combined techniques of retrograde horseradish peroxidase and enkephalin immunocytochemistry. J. Neurosci. 4: 2117-2132, 1984.
105. SAKAI, K., TOURET, M., SALVERT, D., LEGER, L. AND JOUVET, M.: Afferent projections to the cat locus coeruleus as visualized by the horseradish peroxidase technique. Brain Res. 119: 21-41, 1977.
106. SANDKÜHLER, J, AND GEBHART, G, F.: Characterization of inhibition of a spinal nociceptive reflex by stimulation medially and laterally in midbrain and medulla in the pentobarbital anesthetized rat. Brain Res. 305:67-76, 1984a.
107. SANDKÜHLER, J. AND GEBHART, G. F.: Relative contribution of the raphe magnus and adjacent medullary reticular formation to the inhibition by stimulation in the periaqueductal gray of a spinal nociceptive reflex in the pentobarbital-anesthetized rat. Brain Res. 305: 77-87, 1984b.
108. SAWYNOK, J. AND REID, A.: Noradrenergic and purinergic involvement in spinal antinociception by 5-hydroxytryptamine and 2-methyl-5-hydroxytryptamine. Eur. J. Pharmacol. 204: 301-309, 1991.
109. SCHRODER, H. D. AND SKAGERBERG, G.: Catecholamine innervation of the caudal spinal cord in the rat. J. Comp. Neurol. 242: 358-368, 1985.
110. SEYBOLD, V. S. AND ELDE, R. P.: Receptor autoradiography in thoracic spinal cord: Correlation of neurotransmitter binding sites with sympathoadrenal neurons. J. Neurosci. 4: 2533-2542, 1984.
111. SKAGERBERG, G. AND BJÖRKLUND, A.: Topographic principles in the spinal projections of serotonergic and non-serotonergic brainstem neurons in the rat. Neuroscience 15: 445-480, 1985.
112. SLUKA, K. A. AND WESTLUND, K. N.: Spinal projections of the locus coeruleus and the nucleus subcoeruleus in the Harlan and the Sasco Sprague-Dawley rat. Brain Res. 579: 67-73, 1992.
113. SLUKA, K. A. AND WESTLUND, K. N.: Centrally administered non-NMDA but not NMDA receptor antagonists block peripheral knee joint inflammation. Pain 55: 217-225, 1993.
114. SLUKA, K. A., WILLIS, W. D. AND WESTLUND, K. N.: Joint inflammation and hyperalgesia are reduced by spinal bicueulline. NeuroReport 5: 109-112, 1993.
115. SMITH, D. J., MONROE, P. J., KIRK, H. R., CRISP, T. AND SMITH, D. L.: Pharmacological evaluation of neuronal mechanisms involved in spinal antinociception may be confounded by a lack of drug specificity: An analysis using opioid and monoaminergic interactions. Analgesia, 2: 31-41, 1996.
116. SOJA, P. J. AND SINCLAIR, J. G.: Tonic descending influences on cat spinal dorsal horn neurons. Somatosens. Res. 1: 83-93, 1983.
117. STEVENS, R. T., HODGE, C. J. AND APKAHIAN, A. V.: Kolliker-Fuse nucleus: The principal source of pontine catecholamine cells projecting to the lumbar spinal cord of cat. Brain Res. 239: 589-594, 1982.
118. SULLIVAN, A. F., DASHWOOD, M. R. AND DICKENSON, A. H.: Alpha-2-adrenoceptor modulation of nociception in rat spinal cord: Location, effects and interaction with morphine. Eur. J. Pharmacol. 138: 169-177, 1987.
119. SWANSON, L. W. AND HARTMAN, B. K.: The central adrenergic system: An immunofluorescence study of the location of cell bodies and their efferent connections in the rat utilizing dopamine-β-hydroxylase as a marker. J. Comp. Neurol. 163: 467-506, 1975.
120. TAKANO, Y. AND YAKSH, T. L.: Relative efficacy of spinal alpha-2 agonists, dexmedetomidine, clonidine and ST-91, determined in vivo by using N-ethoxycarbonyl-2-ethoxy-1,2-dihydroquinoline, an irreversible antagonist. J. Pharmacol. Exp. Ther. 258: 438-446, 1991.
121. TAMSEN, A. AND GORDH, T.: Epidural clonidine produces analgesia. Lancet 2: 231-232, 1984.
122. TODD, A. J. AND MCKENZIE, J.: GABA-immunoreactive neurons in the dorsal horn of the rat spinal cord. Neuroscience 31: 799-806, 1989.
123. TODD, A. J. AND MILLAR, J.: Receptive fields and responses to iontophoretically applied noradrenaline and 5-hydroxytryptamine of units recorded in laminae I-III of cat dorsal horn. Brain Res. 288:159-167, 1983.
124. Tsou, K. AND JANG, C. S.: Studies on the site of analgesia action of morphine by intracerebral microinjection. Sei Sin 13: 1099-1109, 1962.
125. UNNERSTALL, J. R., KOPAJTIC, T. A. AND KUHAR, M. J.: Distribution of alpha-2 agonist binding sites in the rat and human central nervous sytem: Analysis of some functional, anatomic correlates of the pharmacologic effects of clonidine and related adrenergic agents. Brain Res. Rev. 7: 69-101, 1984.
126. VAN BOCKSTAELE, E. J., PIERIBONE, V. A. AND ASTON-JONES, G.: Diverse afferents converge on the nucleus paragigantocellularis in the rat ventrolateral medulla: Retrograde and anterograde tracing studies. J. Comp. Neurol. 290: 561-584, 1989.
127. VILLANUEVA, L., CHITOUR, D. AND LEBARS, D.: Effects of tizanidine (DS 103-282) on dorsal horn convergent neurons in the rat. Pan 35: 187-197, 1988.
128. WALL, P. D.: The laminar organization of dorsal horn and effects of descending impulses. J. Physiol. (Lond.) 188: 403-423, 1967.
129. WEST, W. L., YEOMANS, D. C. AND PROUDFIT, H. K.: The function of noradrenergic neurons in mediating antinociception induced by electrical stimulation of the locus coeruleus in two different sources of Sprague-Dawley rats. Brain Res. 626: 127-135, 1993.
130. WESTLUND, K. N., BOWKER, R. M., ZIEGLER, M. G. AND COULTER, J. D.: Origins of spinal noradrenergic pathways demonstrated by retrograde transport of antibody to dopamine-β-hydroxylase. Neurosci. Lett. 25: 243-249, 1981.
131. WESTLUND, K. N., BOWKER, R. M., ZIEGLER, M. G. AND COULTER, J. D.: Descending noradrenergic projections and their spinal terminations. Prog. Brain Res. 57: 219-238, 1982.
132. WESTLUND, K. N., BOWKER, R. M., ZIEGLER, M. G. AND COULTER, J. D.: Noradrenergic projections to the spinal cord of the rat. Brain Res. 263: 15-31, 1983.
133. WESTLUND, K. N., BOWKER, R. M., ZIEGLER, M. G. AND COULTER, J. D.: Origins and terminations of descending noradrenergic projections to the spinal cord of monkey. Brain Res. 282: 1-16, 1984.
134. WESTLUND, K. N., CARLTON, S. M., ZHANG, D. AND WILLIS, W. D.: Direct catecholaminergic innervation of primate spinothalamic tract neurons. J. Comp. Neurol. 299: 178-186, 1990.
135. WILCOX, G. L., CARLSSON, K.-H., JOCHIM, A. AND JURNA, I.: Mutual potentiation of antinociceptive effects of morphine and clonidine on motor and sensory responses in rat spinal cord. Brain Res. 405: 84-93, 1987.
136. WILCOX, T. K., YEZIERSKI, R. P. AND WILLIS, W. D.: Inhibition of responses of primate spinothalamic tract (STT) neurons by stimulation of periaqueductal gray. In IASP Abstracts, Edinburgh, 1981.
137. WILLCOCKSON, W. S., CHUNG, J. M., HORI, Y., LEE, K. H. AND WILLIS, W. D.: Effects of iontophoretically released amino acids and amines on primate spinothalamic tract cells. J. Neurosci. 4: 732-740, 1984.
138. WILLIS, W. D., HABER, L. H. AND MARTIN, R. F.: Inhibition of spinothalamic tract cells and interneurons by brain stem stimulation in the monkey. J. Neurophysiol. 40: 968-981, 1977.
139. WILLIS, W. D.: Mechanisms of medial brainstem stimulation-induced inhibition in spinothalamic neurons. In Brainstem Control of Spinal Mechanism, ed. by B. H. Sjolund and A. Bjorklund, pp. 411-438, Fernstrom Foundation Series, Elsevier Biomedical, Amsterdam, 1982.
140. WILLIS, W. D.: Anatomy and physiology of descending control of nociceptive responses of dorsal horn neurons: Comprehensive review. Prog. Brain Res. 77: 1-29, 1988.
141. WILLIS, W. D. J. AND COGGESHALL, R. E.: Functional organization of dorsal horn interneurons. In Sensory Mechanisms of the Spinal Cord, ed. by W. D. Willis and R. E. Coggeshall, pp. 153-216, Plenum Press, New York, 1991.
142. YAKSH, T. L.: Pharmacology of spinal adrenergic systems which modulate spinal nociceptive processing. Pharmacol. Biochem. Behav. 22: 845-858, 1985.
143. YAKSH, T. L. AND RUDY, T. A.: Narcotic analgetics: CNS sites and mechanism of action as revealed by intracerebral injection techniques. Pain 4: 299-359, 1978.
144. YOUNG, W. S. I. AND KUHAR, M. J.: Noradrenergic α1 and α2 receptors: Light microscopic and autoradiographic localization. Proc. Natl. Acad. Sci. USA 77: 1696-1700, 1980.