斑马鱼松果体对视觉敏感性生物钟的维持作用及长时记忆缺陷突变体的筛选
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摘要
生物钟在自然界的多种生物,如动物、植物和真菌中普遍存在。从单细胞的分裂生殖到植物的光合作用,动物的繁殖和激素分泌,以及在分子水平上,都有生物节律的存在。昼夜节律是部分原核生物和所有真核生物基本特征。斑马鱼作为一种在复杂性和简单性方面取得了很好平衡的二倍体脊椎动物,因其个体小,繁殖能力强,体外受精和发育,发育周期短,胚胎和幼鱼透明等独特优点成为理想的可应用于大规模遗传研究的脊椎动物模式生物。同时由于斑马鱼参与节律调控的主要分子机制与哺乳动物比较一致,其也成为适合研究生物节律的模式脊椎动物。
在非哺乳脊椎动物中,松果体作为一个中枢起搏器调节动物行为和生理的昼夜节律。松果体感光细胞的生物钟性质,包括视蛋白表达的日夜转换和视网膜敏感性的振荡都已经得到了很好的数据证实。然而,调节视觉敏感性昼夜节律的基本机制和松果体中枢起搏器调节视觉敏感性昼夜节律功能的可能机制仍然有待发现。本研究中,我们构建了一个转基因斑马鱼品系[Tg(Gnat2:ga14-VP16/UAS:nfsB-mCherry)]。在这个转基因品系中,在松果体感光细胞中表达大肠杆菌硝基还原酶NTR。在胚胎早期发育中,转基因在视网膜和松果体的感光细胞中表达。随着年龄增长,在视网膜感光细胞中的转基因表达逐渐消失。直到第八个月,Gnat2启动子驱动的硝基还原酶不再在视网膜感光细胞中表达,但是仍然在松果体感光细胞中稳定表达。这提供了一种能够特异性杀死松果体感光细胞的模型,例如用甲硝唑处理该转基因鱼后,硝基还原酶NTR可以把药物前体甲硝唑转化为有毒DNA链接交联剂引起细胞死亡,从而特异性杀死松果体感光细胞。在松果体感光细胞缺失的情况下,斑马鱼的行为视觉敏感性保持不变,但是视锥细胞视杆细胞敏感性的节律减弱了。短时间的光照可以一定程度的修复这种行为视觉敏感性生物钟节律的减弱程度。综上,这些结果表明视网膜感光细胞能够对外界环境刺激产生反应,而且能产生视觉敏感性的生物钟节律。然而,仅仅是视网膜感光细胞还不足以维持这种节律。松果体感光细胞中产生的细胞信号可能对维持这种视觉敏感性的生物钟节律有一定贡献。
学习和记忆是中枢神经系统高级活动的一种方式,是人类认知的基础。学习是在经验基础上改变行为反应的过程,记忆是学习到信息贮存和读出的过程。记忆根据持续时间的长短分为长时记忆和短时记忆。许多研究聚焦于学习和记忆如何进行的。但是学习和记忆作为一种复杂的行为,仍然有许多问题需要去探索研究。而在脊椎动物中发现学习和记忆的突变基因,对其机制的研究会有一定的贡献。本实验中以斑马鱼为模式动物,经ENU州诱变及大规模遗传筛选,利用抑制逃避反应的行为学方法获得一例学习记忆的斑马鱼突变体fgt。这种fgt斑马鱼突变体的训练后24小时的长时记忆显著的低于野生型。fgt突变体表现出正常的运动活性和正常的对黑色的倾向性。该突变体的F2代在训练后的24小时的长时记忆中有将近一半(13/30)显著的低于野生型,而另一半则相对正常。同时,在对一个新的环境的探索后,与学习记忆相关的即刻早期基因IEGs c-fos在该突变体将近一半F2代中的表达与野生型的对照有显著性的差异(13/30),另外一半相对正常,与行为学结果一致。这些结果表明,该突变体fgt是一个学习和记忆缺陷的显性突变。而该fgt突变体的发现对未来进一步学习和记忆相关的机制和信号通路提供了一种可能。
Circadian clock exist widely in a variety of organisms, such as animals, plants and fugi. Biological rhythm is present in fission of monoplasts, the photosynthesis of plants, breeding and hormone secretion of animals, as well as the molecular level. Circadian rhythm is one of basic characteristics in parts of prokaryotes and all of eukaryotes. Zebrafish (Danio rerio) is a diploid vertebrate with a good balance of complexity and simplicity. Zebrafish with small size, powerful reproductive capacity, embryo in vitro fertilization and development, short cyclogenl and transparent etc., is the ideal modle vertebrates for large-scale genetic research. Meanwhile, zebrafish also becomes a model animal for study in biological rhythm since the molecular mechanism of rhythm regulatory in zebrafish is similar to mammal.
In non-mammalian vertebrates, the pineal gland functions as the central pacemaker that regulates the circadian rhythms of animal behavior and physiology. The circadian nature of pineal photoreceptor cells, the day-night fluctuation of opsin expression and the oscillation of retinal sensitivity have been well documented. However, the fundamental mechanisms that regulate the circadian rhythms of visual sensitivity and the possible involvement of pineal central pacemakers in the regulation of the circadian rhythms of visual system functions remain to be examined. We generated a transgenic zebrafish line [Tg(Gnat2:gal4-VP16/UAS:nfsB-mCherry)] in which the E.coli nitroreductase(NTR) is expressed in pineal photoreceptor cells. In developing embryos and young adults, the transgene is expressed in both retinal and pineal photoreceptor cells. During aging, the expression of the transgene in retinal photoreceptor cells gradually diminishes. By8months of age, the Gnat2promoter-driven nitroreductase is no longer expressed in retinal photoreceptor cells, but its expression in pineal photoreceptor cells persists. This provides a tool for selective ablation of pineal photoreceptor cells, i.e., by treatments in transgenic fishes with metronidazole, the NTR enzyme reduces nitroimidazole prodrugs metronidazole into cytotoxins that generate DNA interstrand crosslinking and specifically induce death of pineal photoreceptor. In the absence of pineal photoreceptor cells, the behavioral visual sensitivity of the fish remains unchanged; however, the circadian rhythms of rod and cone sensitivity are diminished. Brief light exposures restore the circadian rhythms of behavioral visual sensitivity.Together, the data suggest that retinal photoreceptor cells respond to environmental cues and are capable of entraining the circadian rhythms of visual sensitivity; however, they are insufficient for maintaining the rhythms. Cellular signals from the pineal photoreceptor cells may be required for maintaining the circadian rhythms of visual sensitivity。
Learning and memory, the basis of human cognitive, are mode of advanced activity of central nervous system. Learning is the ability to modify behavioral responses dependent on experience and memory is retention of this change for variable lengths of time, they characterize all animal species. Memory formation further into two distinct phases:short-term memory, and long-term memory. Studies focus on how learning and memory works. However, there are still many mysteries waiting for us to explore in learning and memory as a kind of complex behavior. Finding mutant genes in learning and memory contribute to the study of the mechanism in vertebrate animals. In this study we use zebrafish as a mode animal, isolate a zebrafish mutant in learn and memory named fgt by inhibitory avoidance assay in ENU mutagenesis and large-scale genetic screen. Long-term memory is weak in24hr after training in this fgt zebrafish mutant. The mutant shows normal locomotor activity and normal sensitivity to dark preference. About half of fgt mutant F2(13/30) have bad long-term memory when24hr after training. The expression of IEGs c-fos in half of fgt mutant F2(13/30) after learning in a novel environment increase distinctly from the expression in wild-type fishes, which were in accordance with the behavioral results. These data show that the mutant fgt is a dominant mutant with defect in long-term learning and memory. The identification of fgt mutant may provide a possibility in mechanism and signal pathway of learning and memory.
在非哺乳脊椎动物中,松果体作为一个中枢起搏器调节动物行为和生理的昼夜节律。松果体感光细胞的生物钟性质,包括视蛋白表达的日夜转换和视网膜敏感性的振荡都已经得到了很好的数据证实。然而,调节视觉敏感性昼夜节律的基本机制和松果体中枢起搏器调节视觉敏感性昼夜节律功能的可能机制仍然有待发现。本研究中,我们构建了一个转基因斑马鱼品系[Tg(Gnat2:ga14-VP16/UAS:nfsB-mCherry)]。在这个转基因品系中,在松果体感光细胞中表达大肠杆菌硝基还原酶NTR。在胚胎早期发育中,转基因在视网膜和松果体的感光细胞中表达。随着年龄增长,在视网膜感光细胞中的转基因表达逐渐消失。直到第八个月,Gnat2启动子驱动的硝基还原酶不再在视网膜感光细胞中表达,但是仍然在松果体感光细胞中稳定表达。这提供了一种能够特异性杀死松果体感光细胞的模型,例如用甲硝唑处理该转基因鱼后,硝基还原酶NTR可以把药物前体甲硝唑转化为有毒DNA链接交联剂引起细胞死亡,从而特异性杀死松果体感光细胞。在松果体感光细胞缺失的情况下,斑马鱼的行为视觉敏感性保持不变,但是视锥细胞视杆细胞敏感性的节律减弱了。短时间的光照可以一定程度的修复这种行为视觉敏感性生物钟节律的减弱程度。综上,这些结果表明视网膜感光细胞能够对外界环境刺激产生反应,而且能产生视觉敏感性的生物钟节律。然而,仅仅是视网膜感光细胞还不足以维持这种节律。松果体感光细胞中产生的细胞信号可能对维持这种视觉敏感性的生物钟节律有一定贡献。
学习和记忆是中枢神经系统高级活动的一种方式,是人类认知的基础。学习是在经验基础上改变行为反应的过程,记忆是学习到信息贮存和读出的过程。记忆根据持续时间的长短分为长时记忆和短时记忆。许多研究聚焦于学习和记忆如何进行的。但是学习和记忆作为一种复杂的行为,仍然有许多问题需要去探索研究。而在脊椎动物中发现学习和记忆的突变基因,对其机制的研究会有一定的贡献。本实验中以斑马鱼为模式动物,经ENU州诱变及大规模遗传筛选,利用抑制逃避反应的行为学方法获得一例学习记忆的斑马鱼突变体fgt。这种fgt斑马鱼突变体的训练后24小时的长时记忆显著的低于野生型。fgt突变体表现出正常的运动活性和正常的对黑色的倾向性。该突变体的F2代在训练后的24小时的长时记忆中有将近一半(13/30)显著的低于野生型,而另一半则相对正常。同时,在对一个新的环境的探索后,与学习记忆相关的即刻早期基因IEGs c-fos在该突变体将近一半F2代中的表达与野生型的对照有显著性的差异(13/30),另外一半相对正常,与行为学结果一致。这些结果表明,该突变体fgt是一个学习和记忆缺陷的显性突变。而该fgt突变体的发现对未来进一步学习和记忆相关的机制和信号通路提供了一种可能。
Circadian clock exist widely in a variety of organisms, such as animals, plants and fugi. Biological rhythm is present in fission of monoplasts, the photosynthesis of plants, breeding and hormone secretion of animals, as well as the molecular level. Circadian rhythm is one of basic characteristics in parts of prokaryotes and all of eukaryotes. Zebrafish (Danio rerio) is a diploid vertebrate with a good balance of complexity and simplicity. Zebrafish with small size, powerful reproductive capacity, embryo in vitro fertilization and development, short cyclogenl and transparent etc., is the ideal modle vertebrates for large-scale genetic research. Meanwhile, zebrafish also becomes a model animal for study in biological rhythm since the molecular mechanism of rhythm regulatory in zebrafish is similar to mammal.
In non-mammalian vertebrates, the pineal gland functions as the central pacemaker that regulates the circadian rhythms of animal behavior and physiology. The circadian nature of pineal photoreceptor cells, the day-night fluctuation of opsin expression and the oscillation of retinal sensitivity have been well documented. However, the fundamental mechanisms that regulate the circadian rhythms of visual sensitivity and the possible involvement of pineal central pacemakers in the regulation of the circadian rhythms of visual system functions remain to be examined. We generated a transgenic zebrafish line [Tg(Gnat2:gal4-VP16/UAS:nfsB-mCherry)] in which the E.coli nitroreductase(NTR) is expressed in pineal photoreceptor cells. In developing embryos and young adults, the transgene is expressed in both retinal and pineal photoreceptor cells. During aging, the expression of the transgene in retinal photoreceptor cells gradually diminishes. By8months of age, the Gnat2promoter-driven nitroreductase is no longer expressed in retinal photoreceptor cells, but its expression in pineal photoreceptor cells persists. This provides a tool for selective ablation of pineal photoreceptor cells, i.e., by treatments in transgenic fishes with metronidazole, the NTR enzyme reduces nitroimidazole prodrugs metronidazole into cytotoxins that generate DNA interstrand crosslinking and specifically induce death of pineal photoreceptor. In the absence of pineal photoreceptor cells, the behavioral visual sensitivity of the fish remains unchanged; however, the circadian rhythms of rod and cone sensitivity are diminished. Brief light exposures restore the circadian rhythms of behavioral visual sensitivity.Together, the data suggest that retinal photoreceptor cells respond to environmental cues and are capable of entraining the circadian rhythms of visual sensitivity; however, they are insufficient for maintaining the rhythms. Cellular signals from the pineal photoreceptor cells may be required for maintaining the circadian rhythms of visual sensitivity。
Learning and memory, the basis of human cognitive, are mode of advanced activity of central nervous system. Learning is the ability to modify behavioral responses dependent on experience and memory is retention of this change for variable lengths of time, they characterize all animal species. Memory formation further into two distinct phases:short-term memory, and long-term memory. Studies focus on how learning and memory works. However, there are still many mysteries waiting for us to explore in learning and memory as a kind of complex behavior. Finding mutant genes in learning and memory contribute to the study of the mechanism in vertebrate animals. In this study we use zebrafish as a mode animal, isolate a zebrafish mutant in learn and memory named fgt by inhibitory avoidance assay in ENU mutagenesis and large-scale genetic screen. Long-term memory is weak in24hr after training in this fgt zebrafish mutant. The mutant shows normal locomotor activity and normal sensitivity to dark preference. About half of fgt mutant F2(13/30) have bad long-term memory when24hr after training. The expression of IEGs c-fos in half of fgt mutant F2(13/30) after learning in a novel environment increase distinctly from the expression in wild-type fishes, which were in accordance with the behavioral results. These data show that the mutant fgt is a dominant mutant with defect in long-term learning and memory. The identification of fgt mutant may provide a possibility in mechanism and signal pathway of learning and memory.
引文
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