青海沙蜥成体神经发生的特征描述
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摘要
成体神经发生(adult neurogenesis)是指在成年动物的中枢神经系统中某些特定部位的前体细胞终生不断产生神经元和胶质细胞,新产生的神经元能发育分化为功能性神经元并与其它已经存在的神经元形成突触联系,融合到原有的神经通路中。成体神经发生在物种之间表现出很大差异性。在过去的十年中,有关成体神经发生的调控和功能的研究取得了显著进步,但是对具有自发再生能力的羊膜动物爬行类来说,成体神经发生所发挥的功能仍然知之甚少。本研究以我国青藏高原典型的爬行动物——青海沙蜥为动物模型,对蜥蜴中枢神经系统成体神经发生的基本规律和特征进行观察和研究。利用BrdU活体注射标记分裂细胞,应用免疫组化DAB显色法和免疫组化荧光标记法结合激光共聚焦显微照相技术观察和分析青海沙蜥端脑成体神经干细胞的分布部位和增殖速率。利用经典高尔基染色法描述了青海沙蜥内皮质中主要神经细胞的形态。DCX和GFAP免疫荧光染色分别显示蜥蜴大脑新生神经元和胶质细胞系的总体模式。研究结果表明：
1.与哺乳动物相比,青海沙蜥的大脑皮层构筑较简单,呈典型的三层结构：一条密集的细胞带被两侧的丛状层夹在中间。蜥蜴端脑不具有海马结构。主嗅球和副嗅球都不呈典型的层状结构,细胞排列不规则。
2.最后一次注射BrdU24小时后,标记的新生细胞绝大多数分布于侧脑室的室带,但其它端脑分区中也分布有少量BrdU阳性细胞,其中前嗅核是细胞增殖活动最丰富的区域。中脑和间脑未见BrdU阳性细胞。
3.蜥蜴端脑的内皮质中主要为投射神经元,它们的胞体形态和突起投射各异,或是伸出多条分枝伸向两侧的丛状层,或是聚集成纤维簇向邻近的背皮质区延伸。
4.DCX免疫组化染色结果表明未成熟神经元不仅广泛分布于端脑,而且一些已经迁移进入端脑内皮质中的内丛状层,它们表现出的正在迁移新生神经元的典型特征,一些新生神经元在细胞带中排成一排。
5.青海沙蜥的端脑、间脑和中脑都含有大量GFAP阳性的放射状结构,包括伸长细胞和放射状胶质细胞。它们细长的放射状突起贯穿整个皮层区域,最终形成足板锚定在软脑膜下区。嗅球中的GFAP阳性的波浪状纤维较粗,它们彼此交织成网状结构。游离的星型胶质细胞只存在于蜥蜴的中脑。
青海沙蜥的成体神经发生有其自身特征,新生细胞的增殖、分布和胶质细胞家族的分布模式与种系发育密切相关,这种种系差异主要取决于不同环境刺激和感觉系统生长之间的协同作用。总之,由于爬行类是首先出现星型胶质细胞成分的脊椎动物群,所以其大脑为我们提供了很好的模型来研究神经元的再生和营养信号以及细胞命运分化。因此,这些发现不仅有利于理解不同季节和生态环境对爬行类成体神经发生的影响,而且能帮助我们阐明哺乳动物成年大脑的内源性神经再生能力和兼容性。
Adult neurogenesis, a process of giving rise to neurons and glia continually from progenitors residing in restricted regions of the adult central nervous system (CNS) throughout life, varies considerably across species. Although remarkable progress has been made over the past decade in unraveling the regulation and function of adult neurogenesis, the role of adult neurogenesis in reptiles, the amniotic vertebrates that possess spontaneous regenerative capacity is less known. In the present study, we used Phrynocephalus vlangalii--typical reptile in Qinghai-Tibetan Plateau of China to investigate the general pattern and characteristics of adult neurogenesis.
Lizards were intraperitoneally injected with Bromodeoxyuridine (BrdU) in order to label dividing cells. Distinct immunocytochemistry were performed:DAB-immunostaining and fluorescence-immunostaining, combined with microscope imaging to observe the location and proliferation of newborn cells. Golgi staining technology was used in order to describe the morphology structure of principal neurons in medial cortex of the lizard. DCX-immunostaining and GFAP-immunostaining were carried out to display the general pattern of immature neurons and glial lineage. Our results showed that:
1. Compared with mammals, the brains of P. vlangalii own simpler cytoarchitectonical patterns, but still characteristic of three-laminated fashion:most somata are packed into a cell layer sandwiched between the inner and outer plexiform layers. The cells both in MOB and in AOB of P. vlangalii are irregularly distributed, instead of having laminar organization.
2. Three days after BrdU injection, BrdU-labeled cells were mostly distributed in the ventricular zone of lateral ventricle with cell proliferation highest in region referred to anterior olfactory nucleus (AON).
3. The majority of neurons populating the medial cortex were projection neurons, displaying various soma morphologies. Their projecting processes either reached adjacent dorsal cortical areas and/or the bilateral septum.
4. Immunostaining against DCX showed immature neurons were not only widely distributed in telencephalon, but also had migrated into inner plexiform layer. Some immature neurons orderly arrayed in a line in cell layer of MC and displayed typical migrating newborn neurons.
5. GFAP-immunostaining in telencephalon, diencephalon and mesencephalon were fundamentally represented by radial glial structures, including tanycytes and radial glial cells. Radial GFAP-positive fibers spreaded throughout the entire cortex and terminated with swollen endfeet anchoring the submeningeal surface. The thicker sinuous fibers in OB appeared to interweave together forming a mesh network. An intriguing finding was the emergence of GFAP-positive, star-shaped cells in the mesencephalon.
In conclusion, the adult neurogenesis in lizard is possessed of its own characteristics, which are closely related with phylogenetic differences dependent on the coordinated interaction between different environmental cues and growth of sensory systems. Lizard brain provides us an excellent access to investigate the neuronal regeneration and neurotrophic signals, and cell fate specification, since reptiles represent the first vertebrate group presenting astrocytes proper. Therefore, these findings may not only have implications for understanding the effect of distinctive seasons and ecological environment on the adult neurogenesis of reptiles, but also could favor the elucidation of potential mechanism regulating the endogenous neurogenetic capacity and compatibility in mammalian adult brain.
1.与哺乳动物相比,青海沙蜥的大脑皮层构筑较简单,呈典型的三层结构：一条密集的细胞带被两侧的丛状层夹在中间。蜥蜴端脑不具有海马结构。主嗅球和副嗅球都不呈典型的层状结构,细胞排列不规则。
2.最后一次注射BrdU24小时后,标记的新生细胞绝大多数分布于侧脑室的室带,但其它端脑分区中也分布有少量BrdU阳性细胞,其中前嗅核是细胞增殖活动最丰富的区域。中脑和间脑未见BrdU阳性细胞。
3.蜥蜴端脑的内皮质中主要为投射神经元,它们的胞体形态和突起投射各异,或是伸出多条分枝伸向两侧的丛状层,或是聚集成纤维簇向邻近的背皮质区延伸。
4.DCX免疫组化染色结果表明未成熟神经元不仅广泛分布于端脑,而且一些已经迁移进入端脑内皮质中的内丛状层,它们表现出的正在迁移新生神经元的典型特征,一些新生神经元在细胞带中排成一排。
5.青海沙蜥的端脑、间脑和中脑都含有大量GFAP阳性的放射状结构,包括伸长细胞和放射状胶质细胞。它们细长的放射状突起贯穿整个皮层区域,最终形成足板锚定在软脑膜下区。嗅球中的GFAP阳性的波浪状纤维较粗,它们彼此交织成网状结构。游离的星型胶质细胞只存在于蜥蜴的中脑。
青海沙蜥的成体神经发生有其自身特征,新生细胞的增殖、分布和胶质细胞家族的分布模式与种系发育密切相关,这种种系差异主要取决于不同环境刺激和感觉系统生长之间的协同作用。总之,由于爬行类是首先出现星型胶质细胞成分的脊椎动物群,所以其大脑为我们提供了很好的模型来研究神经元的再生和营养信号以及细胞命运分化。因此,这些发现不仅有利于理解不同季节和生态环境对爬行类成体神经发生的影响,而且能帮助我们阐明哺乳动物成年大脑的内源性神经再生能力和兼容性。
Adult neurogenesis, a process of giving rise to neurons and glia continually from progenitors residing in restricted regions of the adult central nervous system (CNS) throughout life, varies considerably across species. Although remarkable progress has been made over the past decade in unraveling the regulation and function of adult neurogenesis, the role of adult neurogenesis in reptiles, the amniotic vertebrates that possess spontaneous regenerative capacity is less known. In the present study, we used Phrynocephalus vlangalii--typical reptile in Qinghai-Tibetan Plateau of China to investigate the general pattern and characteristics of adult neurogenesis.
Lizards were intraperitoneally injected with Bromodeoxyuridine (BrdU) in order to label dividing cells. Distinct immunocytochemistry were performed:DAB-immunostaining and fluorescence-immunostaining, combined with microscope imaging to observe the location and proliferation of newborn cells. Golgi staining technology was used in order to describe the morphology structure of principal neurons in medial cortex of the lizard. DCX-immunostaining and GFAP-immunostaining were carried out to display the general pattern of immature neurons and glial lineage. Our results showed that:
1. Compared with mammals, the brains of P. vlangalii own simpler cytoarchitectonical patterns, but still characteristic of three-laminated fashion:most somata are packed into a cell layer sandwiched between the inner and outer plexiform layers. The cells both in MOB and in AOB of P. vlangalii are irregularly distributed, instead of having laminar organization.
2. Three days after BrdU injection, BrdU-labeled cells were mostly distributed in the ventricular zone of lateral ventricle with cell proliferation highest in region referred to anterior olfactory nucleus (AON).
3. The majority of neurons populating the medial cortex were projection neurons, displaying various soma morphologies. Their projecting processes either reached adjacent dorsal cortical areas and/or the bilateral septum.
4. Immunostaining against DCX showed immature neurons were not only widely distributed in telencephalon, but also had migrated into inner plexiform layer. Some immature neurons orderly arrayed in a line in cell layer of MC and displayed typical migrating newborn neurons.
5. GFAP-immunostaining in telencephalon, diencephalon and mesencephalon were fundamentally represented by radial glial structures, including tanycytes and radial glial cells. Radial GFAP-positive fibers spreaded throughout the entire cortex and terminated with swollen endfeet anchoring the submeningeal surface. The thicker sinuous fibers in OB appeared to interweave together forming a mesh network. An intriguing finding was the emergence of GFAP-positive, star-shaped cells in the mesencephalon.
In conclusion, the adult neurogenesis in lizard is possessed of its own characteristics, which are closely related with phylogenetic differences dependent on the coordinated interaction between different environmental cues and growth of sensory systems. Lizard brain provides us an excellent access to investigate the neuronal regeneration and neurotrophic signals, and cell fate specification, since reptiles represent the first vertebrate group presenting astrocytes proper. Therefore, these findings may not only have implications for understanding the effect of distinctive seasons and ecological environment on the adult neurogenesis of reptiles, but also could favor the elucidation of potential mechanism regulating the endogenous neurogenetic capacity and compatibility in mammalian adult brain.
引文
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