羟基脲处理对双斑蟋蕈形体结构和嗅觉联合式学习记忆的影响
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摘要
目的：通过对羟基脲处理组和未处理组双斑蟋脑部形态学变化及行为学改变,用以证明蟋蟀蕈形体在蟋蟀联想式学习记忆中起着非常重要的作用。方法：分别对处理组和未处理双斑蟋脑神经节进行常规石蜡切片,苏木精-伊红染色法染色,用以对比处理组和未处理组双斑蟋脑部的结构变化。分别对两组蟋蟀进行嗅觉联想式学习记忆的行为学研究,观察处理组和未处理组的变化。结果：正常的双斑蟋脑神经节由左右对称的两部分组成,包括前脑,中脑,后脑三大部分。其中前脑主要由蕈形体,中央复合体,和视叶三个部分组成。其中蕈形体左右对称的分布在前脑两侧,中央复合体位于中央,视叶位于前脑的两侧,外接复眼。其中蕈形体分为蕈形体冠和蕈形体根部两大部分,在蕈形体冠的上方存在大量密集排列的Kenyon细胞,蕈形体根部分为柄部,以及相互垂直的α叶与p叶。中央复合体则是分为中央体和脑桥两部分。经过过羟基脲处理组的蟋蟀脑依旧分为前脑,中脑,后脑三部分。但羟基脲处理的双斑蟋的蕈形体变化非常大,其蕈形体的完整形态几乎不再存在,蕈形体冠基本消失,蕈形体冠上面的Kenyon细胞的数量大为减少。蕈形体根部只有少量柄部,互相垂直的α叶与β叶基本消失。脑部其它结构变化不大。
未喂食羟基脲组经过训练后PPI均值由训练前的PPI=39.70,变为训练后两小时,训练后两天和四天的PPI值分别问为61.02、55.45和52.17。同时经过SPSS13.0单因素方差分析得出训练前后差异非常显著。而对喂食羟基脲组经过训练后它对两种气味的选择变化非常大,其PPI均值由训练前的PPI=39.34,变为训练后两小时、训练后两天和四天的PPI值分别问为40.17、39.26和40.42。经过SPSS13.0方差分析分析得出训练前后没有着显著性差异。而对两组蟋蟀没有训练前对两种气味的选择使用SPSS13.0单因素方差分析得出训练前后没有着显著性差异。
结论：行为学实验说明羟基脲处理对训练前蟋蟀对气味的选择没有太大的影响,而对训练后的选择影响非常大。结合解剖学实验结果：羟基脲处理在一定程度上破坏了双斑蟋的蕈形体。说明蕈形体在双斑蟋嗅觉联想式学习记忆中起着非常重要的作用。蕈形体的破坏导致双斑蟋嗅觉联想式学习记忆能力的减退甚至消失,但是对其嗅觉本身影响不大。
Objective:This study tries to identify mushroom bodies play a very important role in learning and memory by comparing morphological changes and behavioral brain changes in Gryllus bimaculatus which have been treated with Hydroxyurea to those of control group.
Methods:Conventional paraffin sectioning and hemotoxylin and eosin (HE) staining were adopted to compare the morphological changes of brain in Gryllus bimaculatus between control group and treatment group. In addition, the behavioral research on the associational memory of olfactory sensation was employed to compare the behavioral changes between control group and treatment group.
Results:The Cerebral ganglion in Gryllus bimaculatus is symmetrical from left to right, it include protocerebrum, deutocerebrum and tritocerebrumo The protocerebrum was composed by mushroom bodies, central complex and optic lobes. A Cerebral ganglion have two mushroom bodies, and the mushroom bodies is symmetrical from left to right.Central complex occupied central of the protocerebrum.Optic lobes amonged both sides of protocerebrum.externally ocular.The mushroom bodies composed by calyx and root.There are many Kenyon cell on the top of the calyx.The root of the mushroom bodies divided peduncle, a-lobe andβ-lobe.The central complex was constituted by central body and protocerebral bridge.The Gryllus bimaculatus who was feeded hydroxyurea, whose mushroom bodies changed very much, the mushroom bodies almost lost, the calyx disappeared, the quantity of Kenyon cell was reduced.There is a little peduncle remained in root, anda-lobe andβ-lobe was disappeared.The other part of the cerebral ganglion was no change.
After training the group were not feded Hydroxyurea, whose PPI from before PPI=39.70 becomes,2h PPI=61.02,2d PPI=55.45 and 4d PPI=52.17.Used SPSS13.0, we find thai it changed significant. After training the group were feded Hydroxyurea, whose PPI from before PPI=39.34 becomes,2h PPI=40.71,2d PPI=39.26and 4d PPI=40.42.Used SPSS13.0, we find thai it changed not significant. Before training comparison the fed and not fed grops we found that it was no change.
Conclusion:Behavior experiment illustration that:Whether fed Hydroxyurea or not to Gryllus bimaculatus can not effect it select odor before training, but that can effect it select odor after training.With anatomical results:hydroxyurea can damaged the mushroom bodies.lt explained that mushroom bodies play an important role.The damage of the mushroom bodies can lead Gryllus bimaculatus lost learning and memory ability about olfactory, but it can not affect its olfactory.
未喂食羟基脲组经过训练后PPI均值由训练前的PPI=39.70,变为训练后两小时,训练后两天和四天的PPI值分别问为61.02、55.45和52.17。同时经过SPSS13.0单因素方差分析得出训练前后差异非常显著。而对喂食羟基脲组经过训练后它对两种气味的选择变化非常大,其PPI均值由训练前的PPI=39.34,变为训练后两小时、训练后两天和四天的PPI值分别问为40.17、39.26和40.42。经过SPSS13.0方差分析分析得出训练前后没有着显著性差异。而对两组蟋蟀没有训练前对两种气味的选择使用SPSS13.0单因素方差分析得出训练前后没有着显著性差异。
结论：行为学实验说明羟基脲处理对训练前蟋蟀对气味的选择没有太大的影响,而对训练后的选择影响非常大。结合解剖学实验结果：羟基脲处理在一定程度上破坏了双斑蟋的蕈形体。说明蕈形体在双斑蟋嗅觉联想式学习记忆中起着非常重要的作用。蕈形体的破坏导致双斑蟋嗅觉联想式学习记忆能力的减退甚至消失,但是对其嗅觉本身影响不大。
Objective:This study tries to identify mushroom bodies play a very important role in learning and memory by comparing morphological changes and behavioral brain changes in Gryllus bimaculatus which have been treated with Hydroxyurea to those of control group.
Methods:Conventional paraffin sectioning and hemotoxylin and eosin (HE) staining were adopted to compare the morphological changes of brain in Gryllus bimaculatus between control group and treatment group. In addition, the behavioral research on the associational memory of olfactory sensation was employed to compare the behavioral changes between control group and treatment group.
Results:The Cerebral ganglion in Gryllus bimaculatus is symmetrical from left to right, it include protocerebrum, deutocerebrum and tritocerebrumo The protocerebrum was composed by mushroom bodies, central complex and optic lobes. A Cerebral ganglion have two mushroom bodies, and the mushroom bodies is symmetrical from left to right.Central complex occupied central of the protocerebrum.Optic lobes amonged both sides of protocerebrum.externally ocular.The mushroom bodies composed by calyx and root.There are many Kenyon cell on the top of the calyx.The root of the mushroom bodies divided peduncle, a-lobe andβ-lobe.The central complex was constituted by central body and protocerebral bridge.The Gryllus bimaculatus who was feeded hydroxyurea, whose mushroom bodies changed very much, the mushroom bodies almost lost, the calyx disappeared, the quantity of Kenyon cell was reduced.There is a little peduncle remained in root, anda-lobe andβ-lobe was disappeared.The other part of the cerebral ganglion was no change.
After training the group were not feded Hydroxyurea, whose PPI from before PPI=39.70 becomes,2h PPI=61.02,2d PPI=55.45 and 4d PPI=52.17.Used SPSS13.0, we find thai it changed significant. After training the group were feded Hydroxyurea, whose PPI from before PPI=39.34 becomes,2h PPI=40.71,2d PPI=39.26and 4d PPI=40.42.Used SPSS13.0, we find thai it changed not significant. Before training comparison the fed and not fed grops we found that it was no change.
Conclusion:Behavior experiment illustration that:Whether fed Hydroxyurea or not to Gryllus bimaculatus can not effect it select odor before training, but that can effect it select odor after training.With anatomical results:hydroxyurea can damaged the mushroom bodies.lt explained that mushroom bodies play an important role.The damage of the mushroom bodies can lead Gryllus bimaculatus lost learning and memory ability about olfactory, but it can not affect its olfactory.
引文
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[2]. Matsumoto,Mizunami M.Lifetime olfactory memory in the cricket Gryllus bimaculatus [J].Comparative Physiology A.2002,18(8):295-299.
[3].郑乐怡,归鸿主编.昆虫分类[M].南京：南京师范大学出版社.1999,263-268.
[4].Strambi C,Cayre M,Sattelle DB,etal.Learning [J] Memory,1998,5(1):78-89
[5].李伟红,暴学祥.昆虫脑中蕈形体功能的研究进展[J].淮海工学院学报,2003,12(4)：55-58
[6]. Dujardin F.Memoire surle systeme nerveux des insects[J].Ann SciNat Zool 1850,14:195-206.
[7]. Wolf R,Wittig T,Liu L.Drosophila mushroom bodies are dispensible for visual,tactile and motor learning[J].Learn Memory 1998,5:166-178
[8]. Laurent G,Naraghi M.Odorant-induced oscillations in themushroom bodies of the locust[J].J Neurosci 1994,14:2993-3004.
[9]. Stopfer M,Bhagavan S.Laurent G:Impaired odourdiscrimination on desynchronization of odour-encoding neuralassemblies[J].Nature 1997, 390:70-74.
[10]. Strausfeld NJ,Li Y.Organization of olfactory and multimodalafferent neurons supplying the calyx and pedunculus of the cockroach mushroom bodies[J].J Comp Neurol 1999,40(9):603-625.
[11]. Heisenberg M,Borst A,Wagner S.Drosophila mushroombody mutants are deficient in olfactory learning[J].J Neurogenet 1985,(2):1-30.
[12]. Belle JS,Heisenberg M.Associative odor learning in Drosophila abolished by chemical ablation of mushroom bodies[J].Science 1994,26(3):692-695.
[13]. Connolly JB, Roberts IJ, Armstrong JD. Associative learning disrupted by impaired Gs signaling in Drosophila mushroom bodies[J].Science 1996,27(4):2104-2107.
[14]. Bitterman ME,Menzel R,Fietz A.Classicalconditioningof proboscis extension in honeybees (Apismellifera) [J].J CompPsychol 1983,9(7):107-119.
[15].Tully T,Quinn WG.Classical conditioning and retention in normaland mutant Drosophila melanogaster[J].J Comp Physiol A 1985,15(7):263-277.
[16].Erber J,Masuhr TH,Menzel R.Localization of short-term memory in the brain of the bee, Apis mellifera[J].Physiol Entomol 1980,5:343-358.
[17]. Hammer M.An identified neuron mediates the unconditioned stimulus in associative olfactory learning in honeybees[J].Nature,1993,36(6):59-63.
[18]. Hammer M,Menzel R.Multiple sites of associative odor learning as revealed by local brain microinjections of octopamine in honeybees [J].Learn Memory 1998,(5):146-156.
[19]. Levin LR,Han PL,Hwang PM.The Drosophila learning and memory gene rutabaga encodes a Ca2+/Calmodulin-responsive adenylyl cyclase[J].Cell 1992,68:479-489.
[20].Zars T,Fischer M,Schulz R.Localization of a short-term memory in Drosophila[J].Science 2000,28(8):672-675.
[21].Feany MB,Quinn WG.A neuropeptide gene defined by the Drosophila memory mutant amnesiac[J].Science 1995,26(8):869-873.
[22].DeZazzo J,Xia S,Christensen J.Developmental expression of an amn(+) transgene rescues the mutant memory defect of amnesiac adults[J]. J Neurosci 1999,19:8740-8746.
[23]. Moore MS,DeZazzo J,Luk AY.Ethanol intoxication in Drosophila:genetic and pharmacological evidence for regulation by the cAMP pathway[J].Cell 1998,9(3):997-1007.
[24]. Waddell S,Armstrong JD,Kitamoto T.The amnesiac gene product is expressed in two neurons in the Drosophila brain that are critical for memory[J].Cell 2000,10(3):928-931
[25]. Grotewiel MS,Beck CD.Integrin-mediated short-term memory in Drosophila[J].Nature 1998,391:455-460.
[26]. Han PL,Levin LR,Reed RR.Preferential expression of the Drosophila rutabaga gene in mushroom bodies,neural centers for learning in insects[J].Neuron 1992,9:619-627.
[27]. Nighorn A,Healy MJ,Davis RL:The cyclic AMP phosphodiesterase encoded by the Drosophila dunce gene is concentrated in the mushroom body neuropil[J].Neuron 1991,6:455-467.
[28]. Skoulakis EM,Kalderon D,Davis RL.Preferential expression in mushroom bodies of the catalytic subunit of protein kinase A and its role in learning and memory[J].Neuron 1993,11:197-208.
[29]. Skoulakis EMC,Davis RL:Olfactory learning deficits in mutants for leonardo,a Drosophila gene encoding a 14-3-3 protein[J].Neuron 1996, 17:931-944。
[30].Guo HF,Tong J,Hannan F.A neurofibromatosis-1-regulated pathway is required for learning in Drosophila[J].Nature 2000,40(3):895-898.
[31]. The I,Hannigan GE,Cowley GS.Rescue of a Drosophila NF1 mutant phenotype by protein kinase A[J].Science 1997,27(6):791-794.
[32]. Joiner M,Griffith LC.Visual input regulates circuit configuration in courtship conditioning of Drosophila melanogaster[J].Learn Memory 2000,7:32-42.
[33]. Joiner M,Griffith LC.Mapping of the anatomical circuit of CaM kinase-dependent courtship conditioning in Drosophila[J].Learn Memory 1999,6:177-192.
[34].McBride SM,Giuliani G,Choi C.Mushroom body ablation impairs short-term memory and long-term memory of courtship conditioning in Drosophila melanogaster[J].Neuron 1999,24:967-977。
[35]. Hall JC.The mating of a fly[J].Science 1994,26(4):1702-1714.
[36].Siegel RW,Hall JC,Gailey DA.Genetic elements of courtship in Drosophila:mosaics and learning mutants[J].Behav Genet 1984,14:383-410.
[37]. Neckameyer WS.Dopamine and mushroom bodies in Drosophila [J].Learn Memory 1998,5:157-165.
[38].Bouton ME.Context,time,and memory retrieval in the interference paradigms of Pavlovian learning[J].Psychol Bull 1993,11(4):80-99
[39]. Heisenberg M,Wolf R.Reafferent control of optomotor yaw torque in Drosophila melanogaster[J].J Comp Physiol A 1988,16(3):373-388.
[40]. Wolf R,Heisenberg M.Basic organization of operant behavior as revealed in Drosophila flight orientation[J].J Comp Physiol A 1991,16(9):699-705
[41]. Liu L,Wolf R,Ernst R.Context generalization in Drosophila visual learning requires the mushroom bodies[J].Nature 1999,400:753-756.
[42]. Wustmann G,Heisenberg M.Behavioral manipulation of retrieval in a spatial memory task for Drosophila melanogaster[J].Learn Memory 1997, 4:328-336.
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