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1 Effects of ibotenic acid lesions to rat's hippocampus CA! on place and cue learning in Morris water maze Kana Okada and Hiroshige Okaichi (Doshisha University) It is known that the multiple injections of ibotenic acid to the hippocampal CA l (Cornu Ammonis 1) field of a rat cause cell loss and spatial learning impairment in the place task of Morris water maze. However, no study seems to have been conducted concerning the cue task of the maze. This study examined (1) whether cell loss in the CA 1 field produced learning deficits in both the place and cue tasks; (2) whether training could remedy the deficit in spatial learning; and (3) whether the order of place training and cue training could change the performance outcome. Thirty-seven rats of Wistar strain were divided into Place-Cue (PC) group that received place training first and then cue training, and Cue-Place (CP) group that received training in the reverse order. Half rats in each group were damaged in the CAl field by ibotenic acid injections. Results indicated that damage to CA1 pyramidal cells led to performance deficits only in the place task throughout training, regardless of the sequence of training. The results also indicated that training in the prior task affected learning in the succeeding task, only at the beginning of training, and the effect is transient. Key words: hippocampal CA (cornu ammonis) 1, spatial learning, cue learning, ibotenic acid. The Japanese Journal of Psychology 2005, Vol. 76, No. 4, pp Matsumoto, & Sugita, 1993; Nunn & Jarrard, 1994), Whishaw, 1989; Hagan & Beaughard, 1990; Nunn & Hodges, 1994; Olsen, Scheel, Moller, & Jensen, 1994; Correspondence concerning this article should be sent to: Kana Okada, Department of Psychology, Faculty of Letters, Doshisha University, Karasuma lmadegawa, Kamigyo-ku, Kyoto , Japan ( sunspace@oct.zaq.ne.jp) Buzsaki, Leon, Baimbridge, & Somogyi, 1990; Hsu & Buzsaki, 1993; Matsuyama, Tsuchiyama, Nakamura,
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4 岡 田 岡 市:水 迷 路 学 習 と海 馬CAIイ 355 ボ テ ン酸損 傷 時 を 主 な 学 習 測 度 と し た CP-Control群 内での無処 置 ラ ッ トと疑 似 損 傷 ラ ッ トの 間 に,場 所 課 題 と手 掛 り 課 題 の い ず れ の 逃 避 潜 時 に お い て も有 意 差 が 見 られ な か っ た た め,両 群 を ま と め てCP-Controlと して取 り扱 っ た 訓 練 順 序 の 条 件 に か か わ ら ず,ま 潜 時 を 条 件 群 ご と にFigure2に 目 で は,CP条 ず,場 所 課題 で の 示 し た 1ブ 件 の 潜 時 はPC条 ロ ック 件 の 潜 時 よ り も短 か っ た また 各 条 件 と もControl群 の 潜 時 の 方 がCA1 群 の 潜 時 よ り短 か った 訓 練 が 進 む に つ れ てPC-CA1 群 とCP-CA1群,PC-Control群 CA1 とCP-Control群 の 潜 時 は そ れ ぞ れ 同 じ程 度 に達 した 各 群 の 場 所 課 題 の 逃 避 潜 時 に つ い て,訓 要 因,ブ 練 順 序 と損 傷 の 有 無 を被 験 体 間 ロ ッ ク を被 験 体 内 要 因 とす る3要 因 の 分 散 分 析 を 行 っ た そ の 結 果,CA1損 ブ ロ ッ ク の 主 効 果,ブ 傷 の 有 無 の 主 効 果, ロ ッ ク 訓 練 の 順 序 の 交 互 作 用 が 有 意 で あ っ た(F(1,33)=11.42,p<.01;F(6, 198)=46.99,p<.01;F(6,198)=8.69,p<.01) 更 に ブ ロ ッ ク ご とに訓 練 順 序 の 単純 主効 果 の 検 定 を行 う と,1ブ ロ ッ ク 目 で の み 有 意 で あ っ た(F(1,33)= 18.72,p<.01) 訓練 順 序 の主 効 果 や そ の他 の交 互 作 用 は有意 で は なか った 手 掛 り課 題 に お け る 各 群 の 潜 時 をFigure た 1ブ SHAM ロ ッ ク 目で は,PC群 短 か っ た が,訓 Figure 1. Micrographs of Cresyl violet-stained vertical brain sections, shown in the extent of the damage to hippocanpal CAl of a rat. CAI: Loss of CAI pyramidal cells from multiple injections of ibotenic acid. SHAM: Intact cells of the same CAI region. よ り も潜 時 が 練 が 進 む に つ れ て4群 と も同 程 度 とな っ た 場 所 課 題 の 場 合 と同 様 の3要 っ た 結 果,ブ 因 の分散 分 析 を行 ロ ッ ク の 主 効 果 と ブ ロ ッ ク 訓 練 順 序 の 交 互 作 用 が 有 意 で あ っ た(F(7,245)=5757,p<-01; F(7,245)=5.56,p<.01) ブ ロ ッ ク ご と に訓 練 順 序 の 単 純 主 効 果 の 検 定 を 行 っ た 結 果,1ブ 有 意 で あ っ た(F(1,36)=13.77,p<.01) 題 で は,損 Figure dard error 2. Mean for the escape place latency task. and ranges of one stan- 3に 示 し は,CP群 Figure dard error ロ ッ ク目の み 手 掛 り課 傷 の 有 無 の 主 効 果 と損 傷 の有 無 が か か わ る 3. Mean for escape the cue task. latency and ranges of one stan-
5 Figure 4. Mean first heading direction and ranges of one standard error for the place task. Figure 5. Mean number of entries to the count area and ranges of one standard error for the probe test.
6 Auer, R. N., Jensen, M. L., & Whishaw, I. Q. (1989). Neurobehavioral deficit due to ischemic brain damage limited to half of the CA l sector of the hippocampus. Journal of Neuroscience, 9, Block, F., & Schwarz, M. (1997). Correlation between hippocampal neuronal damage and spatial learning deficit due to global ischemia. Pharmacology, Biochemistry and Behavior, 56, Cho, Y. H., Beracochea, D., & Jaffard, R. (1992). Differential effects of ibotenate lesions of the CAl subfield of the hippocampus on a function of preoperative training in mice. Psychobiology, 20, Freund, T. F., Buzsaki, G., Leon, A., Baimbridge, K.
7 G., & Somogyi, P. (1990). Relationship of neuronal vulnerability and calcium binding protein immunoreactivity in ischemia. Experimental Brain Research, 83, Hagan, J. J., & Beaughard, M. (1990). The effects of forebrain ischemia on spatial learning. Behavioural Brain Research, 41, Hsu, M., & Buzsaki, G. (1993). Vulnerability of mossy fiber targets in the rat hippocampus to forebrain ischemia. Journal of Neuroscience, 13, Jarrard, L. E. (1983). Selective hippocampal lesions and behavior: Effects of kainic acid lesions on performance of place and cue tasks. Behavioral Neuroscience, 97, Jarrard, L. E. (1989). On the use of ibotenic acid to lesion selectively different components of the hippocampal formation. Journal of Neuroscience Methods, 29, Jarrard, L. E. (1991). Use of ibotenic acid to selectively lesion brain structures. Methods in Neurosciences, 7, Jarrard, L. E. (1995). What does the hippocampus really do? Bahevioural Brain Research, 71, Matsuyama, T., Tsuchiyama, M., Nakamura, H., Matsumoto, M., & Sugita, T. (1993). Hilar somatosatin neurons are more vulnerable to an ischemic event than CA1 pyramidal neurons. Journal of Cerebral Blood Flow and Metabolism, 13, Morris, R. G., Garrud, P., Rawlins, J. N., & O'Keefe, J. (1982). Place navigation impaired in rats with hippocampal lesions. Nature, 297, Nunn, J. A., Gray, J. A., & Hodges, H. (1998/1999). Neurotoxic dorsal CA1 lesions versus 4 VO ischemic lesions: Behavioural comparisons. Behavioural Neurology, 11, Nunn, J. A., & Hodges, H. (1994). Cognitive deficits induced by global cerebral ischaemia: Relationship to brain damage and reversal by transplants. Behavioural Brain Research, 65, Nunn, J. A., & Jarrard, L. E. (1994). Silver impregnation reveals neuronal damage in cingulate cortex following 4 VO ischaemia in the rat. Neuroreport, 5, Nunn, J. A., LePeillet, E., Netto, C. A., Hodges, H., Gray, J. A., & Meldrum, B. S. (1994). Global ischemia: Hippocampal pathology and spatial deficits in the water maze. Behavioural Brain Research, 62, Okaichi, H. (1987). Performance and dominant strategies on place and cue tasks following hippocampal lesions in rats. Psychobiology, 15, Okaichi, H. (2001). Effect of dorsal-striatum lesions and fimbria-fornix lesions on the problem-solving strategies of rats in a shallow water maze. Cognitive Affective and Behavioral Neuroscience, 1, O'Keefe, J., & Nadel, L. (1978). The hippocampus as a cognitive map. Oxford: Oxford University Press. Olsen, G. M., Scheel, K. J., Moller, A., & Jensen, L. H. (1994). Relation of spatial learning of rats in the Morris water maze task to the number of viable CA 1 neurons following four-vessel occlusion. Behavioral Neuroscience, 108, Petito, C. K., Feldmann, E., Pulsinelli, W. A., & Plum, F. (1987). Delayed hippocampal damage in humans following cardiorespiratory arrest. Neurology, 37, Save, E., Poucet, B., Foreman, N., & Buhot, M. C. (1992). Object exploration and reactions to spatial and nonspatial changes in hooded rats following damage to parietal cortex or hippocampal formation. Behavioral Neuroscience, 106, Siesjo, B. K. (1981). Cell damage in the brain: A speculative synthesis. Journal of Cerebral Blood Flow & Metabolism, 1, Stubley-Weatherly, L., Harding, J. W., & Wright, J. W. (1996). Effects of discrete kainic acid-induced hippocampal lesions on spatial and contextual learning and memory in rats. Brain Research, 716, Volpe, B. T., Davis, H. P., Towle, A., & Dunlap, W. P. (1992). Loss of hippocampal CA l pyramidal neurons correlates with memory impairment in rats with ischemic or neurotoxin lesions. Behavioral Neuroscience, 106, Whishaw, I. Q., Rod, M. R., & Auer, R. N. (1994). Behavioural deficits revealed by multiple tests in rats with ischemic damage limited to half of the CA l sector of the hippocampus. Brain Research Bulletin, 34, Yanai, S., Okaichi, Y., & Okaichi, H. (2004). Longterm dietary restriction causes negative effects on cognitive function in rats. Neurobiology of Aging, 25, Zola-Morgan, S., Squire, L. R., & Amaral, D. G. (1986). Human amnesia and the medial temporal region: Enduring memory impairment following a bilateral lesion to field CAl of the hippocampus. Journal of Neuroscience, 6, Zola-Morgan, S., Squire, L. R., Rempel, N. L., Clower, R. P., & Amaral, D. G. (1992). Enduring memory impairment in monkeys after ischemic damage to the hippocampus. Journal of Neuroscience, 12,
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