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1.
Asphyxiation of the spinal cord for periods of 2-4 min leads to block of the monosynaptic pathway. At about the same time this blockage takes place, the afferent action potentials fail to invade the presynaptic terminals. Asphyxiation also interferes with the antidromic invasion of motoneurons, and the failure of the antidromic action potentials to invade the motoneuron dendrites coincides with the time of the disappearance of the orthodromic monosynaptic responses. During reoxygenation, both the presynaptic terminals and the dendrites recover their function, or rather their polarization, in a few seconds and yet synaptic transmission reappears only after several minutes. It is postulated that failure of synaptic transmission during asphyxia is due to depolarization of both the presynaptic terminals and the dendrites of the postsynaptic elements. However, repolarization of these elements during reoxygenation, is not sufficient to reestablish synaptic transmission, but recovery of some unidentified biochemical process is apparently necessary.  相似文献   

2.
Asphyxiation of the spinal cord for periods of 2–4 min leads to block of the monosynaptic pathway. At about the same time this blockage takes place, the afferent action potentials fail to invade the presynaptic terminals. Asphyxiation also interferes with the antidromic invasion of motoneurons, and the failure of the antidromic action potentials to invade the motoneuron dendrites coincides with the time of the disappearance of the orthodromic monosynaptic responses. During reoxygenation, both the presynaptic terminals and the dendrites recover their function, or rather their polarization, in a few seconds and yet synaptic transmission reappears only after several minutes. It is postulated that failure of synaptic transmission during asphyxia is due to depolarization of both the presynaptic terminals and the dendrites of the postsynaptic elements. However, repolarization of these elements during reoxygenation, is not sufficient to reestablish synaptic transmission, but recovery of some unidentified biochemical process is apparently necessary.  相似文献   

3.
Potentials of motoneurons of the lower segments of the spinal cord were recorded with the aid of intracellular microelectrodes in experiments on cats with induced tetanus produced by injection of tetanus toxin (1500–2000 mouse LD50) into the extensor muscles of the left shin. Neither afferent volleys of impulses in cutaneous and muscle nerves, nor antidromic volleys in the corresponding ventral roots, produced IPSPs in motoneurons of the extremity into which toxin was injected. The form both of antidromic peak potentials and of monosynaptic EPSPs in motoneurons in which IPSPs were blocked by tetanus toxin did not differ from the form of corresponding potentials of motoneurons in normal cats. The values of threshold depolarization for peak discharges during synaptic and direct stimulation were equal in tetanus and control motoneurons. Resistance and time constant values of the membrane in "tetanus" motoneurons did not differ from the corresponding values for "control" motoneurons.N. I. Pirogov Second Medical Institute, Moscow. Translated from Neirofiziologiya, Vol. 1, No. 1, pp. 25–34, July–August, 1969.  相似文献   

4.
Monosynaptic reflex response of spinal motoneurons to graded afferent volleys has been studied in natural populations and in a representative sample of individual motoneurons. By analysis of input-response relations certain of the requirements for initiation of reflex discharge have been defined. Initation of motoneuron discharge by monosynaptic afferent excitatory volleys results from the development of transmitter potentiality among members of a pool. Transmitter potentiality is considered to have the following characteristics: 1. It is a function of the number of active excitatory synaptic knobs, the degree to which such knobs are aggregated on the motoneuron soma, and the intensity of action per knob. 2. It has an appreciable spatial decrement and rapid temporal decay. 3. While transmitter potentiality has considerable dependence on number of active excitatory knobs, proximity of such knobs is an important variable. Total activation of a discrete zone does not appear to be necessary for initiation of discharge. In addition to initiation of discharge, volleys in monosynaptic afferent excitatory fibers facilitate response otherwise engendered. Such facilitation depends upon the production of an increment in transmitter potentiality. Facilitator potentiality has the following characteristics: 1. It depends principally on number of active excitatory synaptic knobs and intensity of action per knob. 2. Facilitatory action may result from synchronous activity in knobs interspersed among aggregations of knobs otherwise activated, thus fulfilling spatial requirements for transmitter potentiality. Alternatively a residual facilitation may result from a generalized action. 3. Residual facilitation has a slow temporal decay in comparison with transmitter potentiality.  相似文献   

5.
Postsynaptic potentials of 93 motoneurons of the masseter muscle evoked by stimulation of different branches of the trigeminal nerve were studied. Stimulation of the most excitable afferent fibers of the motor nerve of the masseter muscle evoked monosynaptic EPSPs with a latent period of 1.2–2.0 msec, changing into action potentials when the strength of stimulation was increased. A further increase in the strength of stimulation produced an antidromic action potential in the motoneurons with a latent period of 0.9 msec. In some motoneurons polysynaptic EPSPs and action potentials developed following stimulation of the motor nerve to the masseter muscle. The ascending phase of synaptic and antidromic action potentials was subdivided into IS and SD components, while the descending phase ended with definite depolarization and hyperpolarization after-potentials. Stimulation of cutaneous branches of the trigeminal nerve, and also of the motor nerve of the antagonist muscle (digastric) evoked IPSPs with a latent period of 2.7–3.5 msec in motoneurons of the masseter muscle. These results indicate the existence of functional connections between motoneurons of the masseter muscle and its proprioceptive afferent fibers, and also with proprioceptive afferent fibers of the antagonist muscle and cutaneous afferent fibers.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 1, No. 3, pp. 262–268, November–December, 1969.  相似文献   

6.
We studied the antidromic and synaptic potentials evoked from 32 digastric-muscle motoneurons by stimulation of the motor nerve to this muscle, different branches of the trigeminal nerve, and the mesencephalic trigeminal nucleus. Antidromic potentials appeared after 1.1 msec and lasted about 2.0 msec. Stimulation of the infraorbital, lingual, and inferior alveolar nerves led to development of excitatory postsynaptic potentials (EPSP) and action potentials in the motoneurons. The antidromically and synaptically evoked action potentials of the digastric-nerve motoneurons were characterized by weak after-effects. We were able to record EPSP and action potentials in two of the motoneurons investigated in response to stimulation of the mesencephalic trigeminal nucleus, the latent period being 1.3 msec. This indicates the existence of a polysynaptic connection between the mesencephalic-nucleus neurons and the digastric-muscle motoneurons. Eight digastric-muscle motoneurons exhibited inhibitory postsynaptic potentials (IPSP), which were evoked by activation of the afferent fibers of the antagonistic muscle (m. masseter). The data obtained indicate the presence of reciprocal relationships between the motoneurons of the antagonistic muscles that participate in the act of mastication.A. A. Bogomol'ts Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 3, No. 1, pp. 52–57, January–February, 1971.  相似文献   

7.
Electrical Properties of Hypothalamic Neuroendocrine Cells   总被引:7,自引:1,他引:6       下载免费PDF全文
Goldfish hypothalamic neuroendocrine cells have been investigated with intracellular recordings. The cells showed resting potentials of 50 mv and action potentials up to 117 mv followed by a long lasting and prominent diphasic hyperpolarizing afterpotential. The action potential occurred in two steps indicating sequential invasion. "Total" neuron (input) resistance was measured to be 3.3 x 107 Ω and total neuron time constant was 42 msec. Orthodromic volleys, produced by olfactory tract stimulation, generated graded excitatory postsynaptic potentials. These neuroendocrine cells seem, therefore, to have electrical membrane properties that are similar to those of other central neurons. Antidromic volleys (pituitary stimulation) produced inhibitory post-synaptic potentials whose latency was only slightly longer than that of the antidromic spike indicating the presence of recurrent collaterals. This finding suggests that the concept of the neuroendocrine cell as a neuron whose axon forms contacts only on blood vessels and not on other neurons or effector cells is too restrictive. Perfusion of the gills with dilute (0.3 per cent) sea water produced an inhibition of spontaneous activity. This inhibition is discussed in relation to recent work which demonstrates that goldfish hypothalamic hormones facilitate Na+ influx across the gill membrane.  相似文献   

8.
迷走神经感觉输入诱发的鲫鱼Mauthner细胞胞内电位变化   总被引:5,自引:0,他引:5  
Liu LM  Xie Y  Yu F  Zhang MX 《生理学报》2001,53(4):252-256
实验运用微电极穿刺技术,初步探索了刺激鲫鱼右侧迷走神经在双侧Mauthner(M)细胞胞体诱发的胞内电位变化。结果表明:(1)直接刺激鲫鱼右侧迷走神经,可在同侧或对侧M细胞胞体记录到一种短潜伏期、长持续时间、分级的、复合的突触后电位(postsynaptic potentials,PSPs)。此PSPs表现出明显的强度依从性和频率依赖性。(2)刺激迷走神经诱发的PSPs可使逆向锋电位的幅度降低。(3)肌注士的宁后,PSPs的幅度增高、平均持续时间增加、峰值前移。并且可爆发两个以上的动作电位,上述结果提示:迷走神经到M细胞的通路可能 是由长短不等的神经链群组成的。且此通路中不仅包含有兴奋性成分还包含有抑制性成分,而兴奋和抑制之间的相互关系可能起着调节M细胞兴奋性的作用。  相似文献   

9.
Membrane potentials and action potentials evoked by antidromic and direct stimulation were investigated in motoneurons of the trigeminal nucleus in rats innervating the masseter muscle. This motor nucleus was shown to contain cell populations with high and low membrane potentials. The responses of cells of the first group had shorter latent periods of their antidromic action potentials, a longer spike duration, and a lower amplitude and shorter duration of after-hyperpolarization than responses of cells of the second group, and the input resistance of their membrane also is lower. The bimodal character of distribution of electrophysiological parameters of motoneurons in the trigeminal nucleus indicates that "fast" and "slow" fibers of the masseter muscles may be innervated by different types of nerve cells.N. A. Semashko Moscow Medical Stomatological Institute. Translated from Neirofiziologiya, Vol. 13, No. 3, pp. 270–274, May–June, 1981.  相似文献   

10.
Electrical properties of motoneurons in the spinal cord of rat embryos   总被引:5,自引:0,他引:5  
Electrical properties of immature motoneurons were studied in vitro using isolated segments of spinal cords of rat embryos aged 14-21 days of gestation. Stable resting potentials and evoked synaptic potentials were recorded for more than 9 hr, indicating that motoneurons remain viable for many hours. Motoneurons are electrically excitable at 14 days of gestation and from the onset of excitability the action potentials are Na+-dependent but slow rising long-duration Ca2+-dependent action potentials can be evoked if K+ conductance is reduced. Thus, during embryonic development the regenerative potential inward current is Na+-and Ca2+-dependent. During motoneurons' differentiation there are some changes in their electrical properties: resting membrane potential increases, input resistance decreases, input capacitance increases, threshold for action potential decreases, and maximum rate of rise of action potential increases. Afferent motoneuron contacts are formed at 16-18 days of gestation when excitatory synaptic potentials can first be evoked in response to dorsal root stimulation. The changes in input capacitance and threshold for action potential occur at the onset of functional afferent motoneuron contacts, but it is not known whether these changes are autonomous or are influenced by the newly formed sensory inputs.  相似文献   

11.
新生大鼠离体脊髓薄片侧角中间外侧核细胞的电生理特性   总被引:1,自引:0,他引:1  
祝延  马如纯 《生理学报》1989,41(1):63-69
在新生大鼠离体脊髓薄片的中间外侧核作细胞内记录,研究细胞膜的静态与动态电生理特性。细胞的静息电位(RP)变动于-46—-70mV,膜的输入阻抗为108.3±67.9MΩ(X±SD,下同),时间常数9.9±5.6ms,膜电容138.6±124.2pF。用去极化电流进行细胞内刺激时,大部份细胞(85.4%)能产生高频率连续发放,其余细胞(15.6%)仅产生初始单个发放。胞内直接刺激引起的动作电位(AP)幅度为63.4±9.0mV,时程2.4±0.6ms,阈电位水平在RP基础上去极18.7±6.2mV。大部份细胞的锋电位后存在明显的超极化后电位,其幅度为5.1±2.7mV、持续90±31.8ms。刺激背根可在记录细胞引起EPSP或顺向AP,少数细胞尚出现IPSP。而刺激腹根则可引起逆向AP。  相似文献   

12.
1. Electrical stimulation of the rat's contralateral central amygdaloid (CAm) nucleus or the contralateral frontal cortex markedly augmented the antidromic field potential evoked by stimulation of mylohyoid (Myl) nerve. 2. This facilitation was shown to be due to EPSPs of the mylohyoid-anterior digastric (Myl-Dig) motoneurons. 3. In a few motoneurons, cortical EPSPs had fixed short latencies following high-frequency double stimuli and this is believed to be due to a monosynaptic pathway. 4. The amygdaloid or cortically evoked EPSPs relieved IS-SD blockade in a few motoneurons and also facilitated antidromic discharge in others which did not show any IS or M spike response to the same subthreshold antidromic stimulation. The underlying mechanisms are discussed.  相似文献   

13.
Prominent monosynaptic and disynaptic reflex discharges characterize ipsilateral reflex transmission in the third sacral segment. Convergence upon the motoneurons from the two sides of the body is inhibitory, that through disynaptic paths excitatory. The relative latencies of excitation and inhibition of reflex responses, of excitatory and inhibitory synaptic potentials, and of various aspects of impulse discharge in motoneurons are considered. It is concluded: (1) that a direct (i.e. monosynaptic) action of primary afferent collaterals upon motoneurons is responsible for inhibition of monosynaptic reflex discharge of antagonist motoneurons within a myotatic unit; (2) that the inhibitory postsynaptic potential as described is not the primary agency for monosynaptic reflex inhibition of monosynaptic reflex discharge; (3) that, however, a common causal agent may be responsible for inhibition of reflex discharge and for generation of an inhibitory postsynaptic potential; and (4) that the inhibitory post-synaptic potential may be linked with, or be the agent for, inhibition of soma response.  相似文献   

14.
Synaptic inhibition in an isolated nerve cell   总被引:5,自引:0,他引:5       下载免费PDF全文
Following the preceding studies on the mechanisms of excitation in stretch receptor cells of crayfish, this investigation analyzes inhibitory activity in the synapses formed by two neurons. The cell body of the receptor neuron is located in the periphery and sends dendrites into a fine muscle strand. The dendrites receive innervation through an accessory nerve fiber which has now been established to be inhibitory. There exists a direct peripheral inhibitory control mechanism which can modulate the activity of the stretch receptor. The receptor cell which can be studied in isolation was stimulated by stretch deformation of its dendrites or by antidromic excitation and the effect of inhibitory impulses on its activity was analyzed. Recording was done mainly with intracellular leads inserted into the cell body. 1. Stimulation of the relatively slowly conducting inhibitory nerve fiber either decreases the afferent discharge rate or stops impulses altogether in stretched receptor cells. The inhibitory action is confined to the dendrites and acts on the generator mechanism which is set up by stretch deformation. By restricting depolarization of the dendrites above a certain level, inhibition prevents the generator potential from attaining the "firing level" of the cell. 2. The same inhibitory impulse may set up a postsynaptic polarization or a depolarization, depending on the resting potential level of the cell. The membrane potential at which the inhibitory synaptic potential reverses its polarity, the equilibrium level, may vary in different preparations. The inhibitory potentials increase as the resting potential is displaced in any direction from the inhibitory equilibrium. 3. The inhibitory potentials usually rise to a peak in about 2 msec. and decay in about 30 msec. After repetitive inhibitory stimulation a delayed secondary polarization phase has frequently been seen, prolonging the inhibitory action. Repetitive inhibitory excitation may also be followed by a period of facilitation. Some examples of "direct" excitation by the depolarizing action of inhibitory impulses are described. 4. The interaction between antidromic and inhibitory impulses was studied. The results support previous conclusions (a) that during stretch the dendrites provide a persisting "drive" for the more central portions of the receptor cell, and (b) that antidromic all-or-none impulses do not penetrate into the distal portions of stretch-depolarized dendrites. The "after-potentials" of antidromic impulses are modified by inhibition. 5. Evidence is presented that inhibitory synaptic activity increases the conductance of the dendrites. This effect may occur in the absence of inhibitory potential changes.  相似文献   

15.
The present investigation continues a previous study in which the soma-dendrite system of sensory neurons was excited by stretch deformation of the peripheral dendrite portions. Recording was done with intracellular leads which were inserted into the cell soma while the neuron was activated orthodromically or antidromically. The analysis was also extended to axon conduction. Crayfish, Procambarus alleni (Faxon) and Orconectes virilis (Hagen), were used. 1. The size and time course of action potentials recorded from the soma-dendrite complex vary greatly with the level of the cell's membrane potential. The latter can be changed over a wide range by stretch deformation which sets up a "generator potential" in the distal portions of the dendrites. If a cell is at its resting unstretched equilibrium potential, antidromic stimulation through the axon causes an impulse which normally overshoots the resting potential and decays into an afternegativity of 15 to 20 msec. duration. The postspike negativity is not followed by an appreciable hyperpolarization (positive) phase. If the membrane potential is reduced to a new steady level a postspike positivity appears and increases linearly over a depolarization range of 12 to 20 mv. in various cells. At those levels the firing threshold of the cell for orthodromic discharges is generally reached. 2. The safety factor for conduction between axon and cell soma is reduced under three unrelated conditions, (a) During the recovery period (2 to 3 msec.) immediately following an impulse which has conducted fully over the cell soma, a second impulse may be delayed, may invade the soma partially, or may be blocked completely. (b) If progressive depolarization is produced by stretch, it leads to a reduction of impulse height and eventually to complete block of antidromic soma invasion, resembling cathodal block, (c) In some cells, when the normal membrane potential is within several millivolts of the relaxed resting state, an antidromic impulse may be blocked and may set up within the soma a local potential only. The local potential can sum with a second one or it may sum with potential changes set up in the dendrites, leading to complete invasion of the soma. Such antidromic invasion block can always be relieved by appropriate stretch which shifts the membrane potential out of the "blocking range" nearer to the soma firing level. During the afterpositivity of an impulse in a stretched cell the membrane potential may fall below or near the blocking range. During that period another impulse may be delayed or blocked. 3. Information regarding activity and conduction in dendrites has been obtained indirectly, mainly by analyzing the generator action under various conditions of stretch. The following conclusions have been reached: The large dendrite branches have similar properties to the cell body from which they arise and carry the same kind of impulses. In the finer distal filaments of even lightly depolarized dendrites, however, no axon type all-or-none conduction occurs since the generator potential persists to a varying degree during antidromic invasion of the cell. With the membrane potential at its resting level the dendrite terminals contribute to the prolonged impulse afternegativity of the soma. 4. Action potentials in impaled axons and in cell bodies have been compared. It is thought that normally the over-all duration of axon impulses is shorter. Local activity during reduction of the safety margin for conduction was studied. 5. An analysis was made of high frequency grouped discharges which occasionally arise in cells. They differ in many essential aspects from the regular discharges set up by the generator action. It is proposed that grouped discharges occur only when invasion of dendrites is not synchronous, due to a delay in excitation spread between soma and dendrites. Each impulse in a group is assumed to be caused by an impulse in at least one of the large dendrite branches. Depolarization of dendrites abolishes the grouped activity by facilitating invasion of the large dendrite branches.  相似文献   

16.
Responses from motoneurons were recorded with microelectrodes, from the spinal cords of kitten fetuses and newborn kittens between 40 days' gestation and a few days after birth. As in the adult animal, intracellularly recorded action potentials by either ortho- or antidromic shocks have two components, "A" and "B" or IS and SD. The action potentials of the adult and immature motoneuron differ mainly in the afterpotentials which are absent in the fetal cell in "good" condition. Repeated stimulation or deterioration of the cell resulted, however, in the appearance of depolarizing and hyperpolarizing afterpotentials. No major differences were found in the mode of anti- or orthodromic invasion of the adult and fetal motoneuron, but the degree of invasion of the soma-dendritic complex may be somewhat less in the fetal cells. The ventral root discharge by dorsal root stimulation could be obtained in the fetus 3 weeks before birth. This reflex discharge was concluded to be monosynaptic. Excitatory postsynaptic potentials, probably monosynaptically activated, could be recorded from inside motoneurons by stimulation of dorsal root or peripheral nerves. The most remarkable change during prenatal development was an increase in the speed and efficacy of the excitatory synaptic potentials which showed a marked change during the last weeks of prenatal life.  相似文献   

17.
The electrotonically coupled network of about 100 neuroendocrine caudodorsal cells (CDC) of the freshwater snail Lymnaea stagnalis exhibits three states of excitability with distinct electrophysiological characteristics. Transitions between these states occur spontaneously or can be induced experimentally. The CDC produce an ovulation hormone, and the excitability states are clearly related to the egg-laying cycle of the snail. Two hours before egg laying, the cells enter an active state, which lasts one hour. This phase is characterized by a spontaneous firing pattern, which in preparations can be evoked as an afterdischarge, and during which the hormone is thought to be released. After this, the cells enter an inhibited state in which no other activity than directly stimulus-dependent ortho- and antidromic action potentials can be evoked. This phase lasts till about four hours after egg laying. The subsequent resting state is characterized by facilitation of the responses upon repetitive stimulation of the cells, leading to depolarization of the network and additional action potentials. In this phase, an afterdischarge can be evoked, which brings the cells into the active stage again.  相似文献   

18.
The present study demonstrates the enhanced efficacy of impulse initiation among the hindlimb alpha motoneurons of flexor and extensor origins (n = 35) upon electrical stimulation of the locus coeruleus (LC) in decerebrate cats. When combined with the LC-evoked excitatory postsynaptic potential (EPSP), intracellular hyperpolarization-induced partial and total blocks of antidromic invasion were overcome, resulting in full-spike generation in all cells (n = 21). In three other cells, partial blocks, representing the motoneuron refractoriness resulting from double stimulation at close intervals, were relieved by the concomitant LC-EPSP. When an antidromic volley occurred at a time when the somadendritic (SD) membrane was near threshold, LC stimulation was shown to increase the probability of full-spike initiation as well as to shorten the initial segment (IS)-SD delay, suggesting a coerulospinal enhancement of the safety factor for IS-SD impulse conduction. When coincident with the LC-EPSPs, group Ia EPSPs of flexor and extensor origins were demonstrated to reach the threshold of discharging the cells (n = 4). In those cells exhibiting prominent depolarizing synaptic noise (n = 10), LC stimulation was sufficient to cause the cell to fire action potentials presumably by interacting with concomitant excitatory synaptic drive. The present results advocate that the descending LC excitatory drive has engaged in the action potential initiation process of the alpha motoneuron, facilitating its reaching the firing threshold during concurrent depressed membrane excitability as well as subthreshold converging inputs.  相似文献   

19.
The present study was aimed at determining if inputs from the locus coeruleus (LC) have any effect on repetitive firing of ventral horn motoneurons in cats. In hindlimb flexor and extensor motoneurons stimulated intrasomatically with current below the threshold for repetitive discharges, added LC-evoked excitatory post-synaptic potentials (EPSPs) were consistently found to produce repetitive firing, suggesting a lowering in the repetitive firing threshold attributable to excitatory LC inputs. With those extensor motoneurons showing episodic, self-sustained firing, LC-EPSPs introduced during the quiescent period were capable of starting a continuous discharge with rhythmic frequencies higher than the spontaneous activity. In some of these cells, intracellularly applied hyperpolarizing current was able to stop the spontaneous discharges. Subsequently, LC stimuli were found to reinitiate repetitive discharges, thus counteracting the ongoing suppression of the motoneurons. Quantitative analysis of the single-spike afterhyperpolarization (AHP) indicated a consistent reduction in its amplitude and time course (duration, time-to-peak, half-decay time) for flexor and extensor motoneurons in response to LC conditioning stimuli. Present results suggest an excitatory LC action on the repetitive discharges of cat motoneurons accompanied by a concurrent decrease in the amplitude and time course of the single-spike AHPs.  相似文献   

20.
Interactions of peripheral inputs to the motoneuron of the kitten fetus as young as 3 weeks prenatal were studied by reflex discharge from the ventral root as well as by recording from single motoneurons. Facilitation was found between two synergists in fetuses 1 to 2 weeks before birth. Intracellular recording showed that the facilitation could be explained by summation of excitatory postsynaptic potentials. Inhibition was found between antagonists in the fetuses 2 to 3 weeks before birth and was accompanied by inhibitory postsynaptic potentials. Recurrent inhibition was very powerful in the fetal spinal cord as shown by large motoneuron hyperpolarization by antidromic stimulation. Cells presumed to be "Renshaw cells" and which responded to both ortho- and antidromic stimulation with repetitive firing were shown in the 2 weeks prenatal fetus. These results lead to the conclusion that there is considerable effective synaptic connection of afferent collaterals already established by the later stage of intrauterine life and that this may be achieved independently of external stimuli.  相似文献   

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