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[豚鼠皮层的产前发育及产后变化:自然剥夺实验的微观评估。I. 产前发育]

[Prenatal development and postnatal changes in the guinea pig cortex: microscopic evaluation of a natural deprivation experiment. I. Prenatal development].

作者信息

Schüz A

出版信息

J Hirnforsch. 1981;22(1):93-111.

PMID:7240730
Abstract

The present paper is based on the question, to what extent the cortical structure is determined by genetic factors and how far it is dependent on environmental stimuli. Some deprivation experiment in the literature have supported the assumption tha excitation coming from the sense organs contributes to the formation of synaptic connections in the cortex. This made it possible to invoke the formation of synapses (or dendritic spines) as a substrate of learning processes. Results of experiments on the influence of artificial environments on the formation of synapses have been, however, somewhat contradictory. On this background it was interesting to investigate the cortical development of the guinea-pig, an animal which is highly developed at birth. This percocity separates in time the process of genetically determined development from the changes due to environmental stimuli, which are amply overlapping in altricial animals such as mouse, rat, and cat. A comparison between the cortices of prenatal and adult guinea-pigs showed that the density of dendritic spines has reached adult values already before birth (12/10 micrometers dendritic length before birth, 11,5/10 micrometers in adult animals, fig. 5-8, and 17). The counts have been made on basal dendrites of Golgi-impregnated pyramidal cells in the upper third of the cortex. Also, the difference in the density of synapses on electronmicrographs in animals just before birth (8,9 x 10(8)/mm(3)) and in adult animals (9,4 x 10(8)/mm(3)) was not significant (figs. 13, 14, and 16). The samples have been taken from the second cortical layer. The two areas investigated showed small but significant differences in the time course of spine formation. In both areas the density of spines reached a maximum first and then decreased slightly toward the adult values. However, in the postcallosal area the maximum was reached earlier than in the precallosal area (fig. 9). The decrease in spine density after the maximum, about 18% in both areas, may be partly explained by the growth of dendrites. From the increase in brain volume between birth and adult age and from the density of synapses at different stages, one can conclude that the total number of synapses at birth is about two thirds of that in adult animals. Similarly, the proportion of spines present at birth was at least the same or even higher (fig. 12). Thus, most of the connections in the cortex of the guniea pig are formed without the influence of environmental stimuli. This puts strong doubts on the idea of the formation of synapses or dendritic spines as memory traces. Synapses and spines seem to be the prerequisites of learning rather than the result of it. In part II the question will be examined, if postnatal changes in the cortex of the guinea-pig, especially on spines and synapses, are possible candidates for memory traces.

摘要

本文基于这样的问题

皮层结构在多大程度上由遗传因素决定,又在多大程度上依赖于环境刺激。文献中的一些剥夺实验支持了这样的假设,即来自感觉器官的兴奋有助于皮层中突触连接的形成。这使得将突触(或树突棘)的形成作为学习过程的基础成为可能。然而,关于人工环境对突触形成影响的实验结果有些相互矛盾。在此背景下,研究豚鼠的皮层发育很有意思,豚鼠是一种出生时就高度发育的动物。这种早熟在时间上把遗传决定的发育过程与环境刺激引起的变化区分开来,而在诸如小鼠、大鼠和猫等晚成动物中,这两种变化大量重叠。产前和成年豚鼠皮层的比较表明,树突棘的密度在出生前就已达到成年水平(出生前每10微米树突长度有12个树突棘,成年动物为11.5个/10微米,图5 - 8和17)。计数是在皮层上部三分之一处经高尔基染色的锥体细胞的基底树突上进行的。同样,出生前动物(8.9×10⁸/mm³)和成年动物(9.4×10⁸/mm³)电子显微镜照片上突触密度的差异并不显著(图13、14和16)。样本取自第二皮层。所研究的两个区域在树突棘形成的时间进程上有小但显著的差异。在这两个区域,树突棘密度都先达到最大值,然后朝着成年值略有下降。然而,胼胝体后区比胼胝体前区更早达到最大值(图9)。最大值之后树突棘密度的下降,两个区域约为18%,这可能部分是由树突的生长所解释的。从出生到成年大脑体积的增加以及不同阶段突触的密度可以推断,出生时突触的总数约为成年动物的三分之二。同样,出生时存在的树突棘比例至少相同甚至更高(图12)。因此,豚鼠皮层中的大多数连接是在没有环境刺激影响的情况下形成的。这对将突触或树突棘的形成作为记忆痕迹的观点提出了强烈质疑。突触和树突棘似乎是学习的前提条件,而非学习的结果。在第二部分中,将探讨豚鼠皮层出生后的变化,尤其是树突棘和突触的变化,是否可能是记忆痕迹的候选因素。

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