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1.
Summary Horseradish-peroxidase was used to anterogradely label and thus to trace the growth of corticospinal axons in rats ranging in age from one day to six months. Three to eight HRP-gels were implanted in the left cerebral hemisphere of the cortex. In each spinal cord three levels were studied, the cervical intumescence (C5), the mid-thoracic region (T5) and the lumbar enlargement (L3). The methodology employed for the electron microscopic visualization of HRP has been described previously (Joosten et al. 1987a).The outgrowth of labelled unmyelinated corticospinal tract axons in the rat spinal cord primarily occurs during the first ten postnatal days. The outgrowth of the main weve of these fibres is preceded by a number of pathfinding axons, characterized by dilatations at their distal ends, the growth cones. By contrast, later appearing unmyelinated axons, which presumably grow along the pathfinding axons, do not exhibit such growth cones. The first labelled pioneer axons can be observed in the cervical intumescence at postnatal day one (P1), in the mid-thoracic region at day three (P3) and in the lumbar enlargement at day five (P5).Prior to the entrance of the axons, the prospective corticospinal area or the pre-arrival zone is composed of fascicles consisting of unlabelled, unmyelinated fibres surrounded by lucent amorphous structures. During the outgrowth phase of the corticospinal fibres some myelinated axons could be observed within the outgrowth area even before day 14. These axons, however, were never labelled. These findings strongly suggest that the outgrowth area, which is generally denoted as the pyramidal tract, contains other axons besides the corticospinal fibres (and glial cells). The process of myelination of the labelled corticospinal tract axons in the rat spinal cord starts rostrally (C5) at about day 14 and progresses caudally during the third and fourth postnatal weeks. Although myelination seems to be largely complete at day 28 at all three spinal cord levels, some labelled unmyelinated axons are still present in the adult stage.  相似文献   

2.
The growth of the corticospinal tract was studied in prenatal and neonatal rats using the anterograde transport of horseradish peroxidase injected into the cerebral cortex as a marker in lightmicroscopic preparations. The findings were compared with electron-microscopic observations on normal material at the same ages. Labelled corticofugal axons traverse the diencephalon by gestational day 17.5, reach the pontine nuclei by gestational day 19.5, and the caudal limit of the medulla oblongata by gestational day 20.5, just before birth. On the day after birth, labelled corticospinal axons have crossed in the pyramidal decussation and extended into the dorsal columns of the upper cervical spinal cord. Corticospinal axons reach the thoracic segments by postnatal day 3, the lumbar segments by day 6 and the sacral segments by day 9. The lower end of the spinal cord is reached only after postnatal day 14. Beside the principal corticospinal tract in the dorsal columns, two other smaller corticospinal tracts occupy an intermediate position in the base of the cervical dorsal horn and a lateral position in the lateral white column. The intermediate tract is not found below cervical levels. Growth cones are seen at the tips of axons in light- and electron-microscopic material. The first corticospinal axons, less than 0.5 microns in diameter and grouped in tight fascicles, grow through a dense fabric of astrocytic and other glial processes in which no obvious pre-existing channels could be identified. Growth of corticospinal axons into the dorsal horn adjacent to the main tract is delayed until 2–3 days after the initial arrival of the tract at a given segment. This begins in the cervical segments only after the thalamocortical fibers have invaded the sensory-motor cortex though the parent pyramidal cells of the tract are still highly immature. The rate of extension of corticospinal axons is not constant. Growth down the dorsal columns is characterized by accelerated growth spurts on postnatal days 4 and 9. Much slower growth characterizes initial outgrowth through the diencephalon and later ingrowth into the spinal gray matter. There is approximately a three-fold increase in the numbers of corticospinal axons in the cervical segments between postnatal days 5 and 10. Myelination commences between postnatal days 10 and 12.It is concluded that the development of the corticospinal tract in the rat displays features that are common to other developing pathways in the rat and other species. Initial outgrowth of corticospinal axons is independent of afferent innervation, occurring at a time when the parent cell bodies are very immature. The early growth of corticospinal axons to the vicinity of their targets is followed by a substantial waiting period, comparable to that seen in other systems, before final invasion of the target. The factors responsible for the initiation of the second growth spurt, carrying axons into the target gray matter are not known. However, the final invasion of gray matter takes place only after the cells of origin of corticospinal axons have received a substantial afferent input. The rate of initial growth of corticospinal axons down the dorsal columns is not constant, but varies from region to region. Electronmicroscopy has failed to detect any morphological evidence for factors that might guide or promote the growth of corticospinal axons. The majority of corticospinal axons exclusive of the first ‘pathfinders’ seem to grow as tight fascicles in which individual axons contact only one another.  相似文献   

3.
In the present investigation the role of astrocytes and their precursors in guidance of outgrowing corticospinal tract axons in the rat is studied. Antibodies against glial fibrillary acidic protein and vimentin are used to analyse immunogen expression of glial cells, whereas the postnatal outgrowth of corticospinal tract axons through the spinal cord was studied using anterogradely transported horseradish peroxidase. The first, leading corticospinal tract axons, being the objective of the present study, are characterized by dilatations at their distal ends, the growth cones. Growth cones of pioneer corticospinal tract axons are randomly distributed in the presumptive corticospinal tract area of the ventral most part of the dorsal funiculus. A dramatic change in glial cell labelling is found from the majority being vimentin immunoreactive and glial fibrillary acidic protein-negative at birth to almost all being the reverse at the end of the fourth postnatal week. From double labelling experiments it can be concluded that the vimentin-glial fibrillary acidic protein transition occurs within astrocyte precursor cells. The absence of glial fibrillary acidic protein-immunoreactive glial cells during the outgrowth period of pioneer corticospinal tract axons indicates that they cannot play a role in the guidance of outgrowing corticospinal tract pioneer axons. Vimentin-immunoreactive glial cells are present throughout the presumptive corticospinal tract area at the time of arrival of the leading corticospinal tract fibres. The vimentin-immunoreactive glial cells, which themselves are orientated perpendicular to the outgrowing corticospinal tract axons, are mainly arranged in longitudinal tiers parallel to the rostrocaudal axis. Electron microscopically, growth cones of pioneer corticospinal tract axons frequently exhibit protrusions into vimentin-immunoreactive glial cell processes, suggesting an adhesive type of contact. Therefore, in addition to a positional role, vimentin-immunoreactive glial cells probably play a chemical role in guidance of pioneer corticospinal tract axons. A prominent vimentin-immunoreactive glial septum was noted during corticospinal tract outgrowth in the midline raphe of the medulla oblongata and spinal cord whereas it is absent in the decussation area of corticospinal tract fibres. After the first postnatal week the major vimentin-immunoreactive glial barrier either completely disappears (medullary levels) or gradually reduces to a minor glial fibrillary acidic protein-immunoreactive one (spinal cord levels).(ABSTRACT TRUNCATED AT 400 WORDS)  相似文献   

4.
Summary A quantitative electron microscopic analysis was undertaken of the development of the pyramidal tract, at the level of the third cervical spinal segment, in rats ranging in age from the day of birth to three months old. The axon number was calculated as the product of axon density, determined in a systematic random sample of electron micrographs, and tract area. During the first postnatal week the tract contains thin unmyelinated axons and growth cones. Growth cones are abundant in neonatal rats, but can still be observed occasionally at the end of the first postnatal week, indicating a continuous addition of pyramidal tract axons during the first postnatal week. Myelination starts around P10. By the end of the first postnatal month approximately 50% of the axons have already been myelinated. Myelination proceeds during further maturation, but in the three month old rat 28% of the axons are still unmyelinated. The total number of axons increases rapidly after birth up to 153 000 at the fourth postnatal day. Subsequently, the number of axons is reduced by nearly 50% to 79 000 in the adult rat. The axon loss is most prominent during the second postnatal week, when 32 000 axons are climinated, but continues for several weeks at a slower rate.  相似文献   

5.
Summary An anterograde tracer study has been made of the developing corticospinal tract (CST) in the rat using wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP). Analysis of normal Rager stained material revealed that corticospinal axons reach upper cervical spinal cord levels at the day of birth (PO). Postnatal rats ranging in age from one (P1) to fourteen (P14) days received multiple WGA-HRP injections into the cortex of their left hemisphere and were allowed to survive for 24 h. The first labeled CST fibers caudally extend into the third thoracic spinal cord segment at P1; into the eighth thoracic segment at P3; into the first or second lumbar segment at P7 and into the second to third sacral segment at Pg. Thus the outgrowth of the leading pioneer fibers of the CST is completed at P9 but later developing axons are continuously added even beyond P9. Quantitative analysis of the amount of label along the length of the outgrowing CST revealed a characteristic pattern of labeling varying with age. The most striking features of that pattern are: (1) the formation of two standing peaks at the level of the cervical and lumbar enlargements respectively and (2) the transient presence of a smaller running peak which moves caudally with the front of the outgrowing bundle. The standing peaks are ascribed to the branching of the axon terminals at both intumescences, whereas the running peak probably arises by the accumulation of tracer within the growth cones at the tips of the outgrowing CST axons. Factors such as the number of axons, the varying axon diameters, the branching collaterals, the presence of varicosities, the transport rate of the tracer, the uptake of the tracer at the injection site, which possibly may affect the amount of label present in both the entire bundle and in the individual axons are discussed. Current research is focused upon an analysis of the relation between the site of injection within the cortex and the pattern of labeling of the CST. A delay of two days was found between the arrival of the CST axons at a particular spinal cord level and their outgrowth into the adjacent spinal gray. However, combined HRP and electronmicroscopic experiments are necessary to determine the factors behind the maturation of the CST as well as the maturation of the spinal gray.  相似文献   

6.
L1 is a representative of a family of carbohydrate neural cell adhesion molecules. The expression of L1 was studied during postnatal development of the rat pyramidal tract by immunohistology using polyclonal antibodies to L1 in spinal cord cervical intumescences. On postnatal day 1 (P1), L1 immunoreactivity was present in the entire dorsal funiculus, consisting of the ascending fasciculus gracilis and fasciculus cuneatus and the descending pyramidal tract. At that time the cervical pyramidal tract contains the first outgrowing corticospinal axons. At P4 both the fasciculus gracilis and the pyramidal tract are immunoreactive whereas the fasciculus cuneatus is negative. At P10 the pyramidal tract is intensely labelled whereas both ascending bundles are negatively stained. In the period between P4 and P10 the pyramidal tract is characterized by a massive outgrowth of corticospinal axons. During pyramidal tract myelination, between P10 and the end of the third postnatal week (P21), L1 immunoreactivity is progressively reduced. These observations suggest that L1 may play a prominent role in outgrowth, fasciculation and the onset of myelination of rat pyramidal tract axons. The differential L1 immunoreactivity of the pyramidal tract and the earlier developing ascending systems in rat dorsal funiculus indicate that this polyclonal antiserum is a useful differentiating marker for outgrowing fibre tracts.  相似文献   

7.
Anterograde staining with Phaseolus vulgaris leucoagglutinin (PHA-L) revealed the spinal arborization pattern of corticospinal tract (CST) fibers in the cervical enlargement of the rat. Within the confines of the pyramidal tract local nets of small fibers are present in addition to the rather large CST fibers with varicosities. CST termination is primarily located in lamina IV and extends into lamina V and VI. Extensive collateralization of CST axons was found interconnecting neurons located both in different horizontal laminae and in subsequent spinal cord segments. This complex pattern of CST collateralization is suggested to add a coordinative role in motor control to this tract both through serial axo-dendritic contacts in the spinal gray and through axo-axonal contacts in the white as well as the gray matter.  相似文献   

8.
Mutations in the gene encoding the neural recognition molecule L1 result in hypoplasia of the corticospinal tract and path finding errors of corticospinal axons at the pyramidal decussation. Candidate molecules that have been implicated in L1-dependent guidance of corticospinal axons from the ventral medullary pyramids to the contralateral dorsal columns of the cervical spinal cord include Semaphorin3A and CD24. In the present study, we anterogradely labeled corticospinal axons from the sensorimotor cortex of young postnatal Semaphorin3A- and CD24-deficient mice to elucidate potential functions of both proteins during formation of this long axon projection. Our results indicate that elongation, collateralization, fasciculation and path finding of corticospinal axons in mice proceed normally in the absence of Semaphorin3A or CD24.  相似文献   

9.
目的 :为了探索神经特异型基因c src产物 pp60c src( ) 在突触形成中的作用机制。方法 :用免疫细胞化学方法检测pp60c src( ) 在初代培养鸡胚脊神经节细胞体、突起和生长端内的分布特征。结果 :培养 2 4小时的生长端丝状伪足不断运动改变生长端的形态 ,pp60c src( ) 的免疫反应活性分布在神经细胞的核周体、神经突起、生长端部分领域和丝状伪足 ;培养 4 8小时的生长端接近靶细胞 ,丝状伪足的活泼性和运动性明显减弱以至消失 ,在伸长的突起上有pp60c src( ) 免疫反应活性较强的串珠样膨体 ,在生长端体部的部分区域和丝状伪足免疫反应活性明显减弱。结论 :( 1 )在生长端向突触前部分化的过程中 ,pp60c src( ) 免疫活性曾发生一过性的减低或消失 ,它具有时空特异性 ;( 2 )pp60c src( ) 对突触发育具有重要的调控作用。  相似文献   

10.
Retinal projections to the nucleus of the optic tract (NOT) and accessory optic nuclei (AON) were studied in the postnatal hooded rat after monocular injection of cholera toxin B subunit (CTB) into the vitreous chamber of the eye. At all postnatal ages, retinal axons were labeled sensitively; they revealed dense projections to the contralateral, and sparse but distinct projections to the ipsilateral, NOT and AON. The CTB labeling enabled the first delineation of the complete morphology of developing retinal axons in the ipsilateral NOT and AON. From postnatal day (P) 1 to P3, axons with complex growth cones were seen, and unbranched collaterals with simple growth cones increased and extended gradually. At P6, complex growth cones disappeared while branched collaterals with simple growth cones as well as small-sized varicosities increased. By P12 (two days before eye-opening) the adult-like pattern of terminal arbors appeared. The branched collaterals with tiny, small-sized varicosities present probably represented developing synaptic boutons. At P16 (after eye opening), the pattern of terminal arbors was well developed, almost to the same extent as in the adult. By contrast, a broadly distributed, transient retinal projection around NOT and AON was gradually eliminated; it started to disappear during the first few postnatal days, and was fully retracted by the time of eye-opening time to a pattern normal for the adult.  相似文献   

11.
Growing corticospinal axons by-pass lesions of neonatal rat spinal cord   总被引:1,自引:0,他引:1  
The anterograde transport of horseradish peroxidase was used to label newly growing corticospinal axons after they had entered lesioned regions of the neonatal rat spinal cord. Two types of lesions were made at thoracic and lumbar levels before the arrival of the first corticospinal axons. (1) Thermal lesions were produced by the brief application of a heated rod to the vertebral column and could destroy the normal growth path over several spinal segments. Corticospinal axons, when successful in growing distal to thermal lesions, did so at the same rate as in normal controls and retaining their normal relative positions and morphology, especially fasciculation. (2) Surgical lesions were produced by cutting the spinal cord and were limited to one segment but could result in a barrier in the normal growth path composed of a cyst or glial scar. Corticospinal axons that succeeded in growing distal to a surgical lesion did so by being deflected to unusual positions, became defasciculated, and sometimes their normal growth rate was slowed. That corticospinal axons could in many instances grow past the two types lf lesion suggests that a morphologically stereotyped glial scaffolding is not necessary for axon growth. The role of fasciculation in normal axon growth is highlighted by the disparate effects of the two types of lesion.  相似文献   

12.
Summary The differentiation of dendrites in the medial trapezoid nucleus of the opossum and cat has been traced from the stage of the post-migratory neuroblast in developmental series prepared with the rapid Golgi technique. The post-migratory neuroblast is an elongated cell. Its perikaryon is located initially at the outer limiting layer of the medulla. Its primitive internal process grows into the primordial medial trapezoid nucleus and gives rise to an axon. On the part of the neuroblast adjacent to the axon's origin the endings of the afferent axons beging to differentiate. The perikaryon moves to the same part of the neuroblast through the primitive internal process. Subsequently the dendrites differentiate. Dendrites and their branches form from budding growth cones. The cell body and dendritic processes of the young growing neuron are covered with transitory filopodia. Sprouting growth cones and filopodia appear at the tips and along the shafts of the elongating and enlarging dendrites. The locomotor and synthetic activities of the growth cones establish the stereotyped dendritic branching patterns of each kind of neuron. The development of the dendritic branches accompanies the elaboration of the particular type of axonal plexus that will become synaptically related. This suggests that the patterns of the dendritic trees and of the afferent axonal end-branches derive from mutual interactions of the growing dendritic and axonal branches. These interactions may be mediated by physical contacts as well as chemotactic factors. The filopodia are implicated in the formation of dendritic appendages. Filopodia could participate in membrane synthesis, locomotion, and synaptogenesis. There is an indication that the afferent axons can induce the differentiation of the post-synaptic parts of the neuroblast. The findings imply that the influence of physical and chemical factors in the differentiation of the synaptic organization of the brain depends on their temporal and spatial sequences.Supported in part by U.S. Public Health Service Research Grant NB 06115 to Harvard University.With the technical aid of Mrs. R. R. Morest and of Miss P. E. Palmer.  相似文献   

13.
The mammalian corticospinal tract is known to contain axons that travel from the cerebral cortex to various levels of the spinal cord and its main function is thought to be the mediation of voluntary movement. The current study describes neuroanatomy related to the corticospinal tract of the giraffe. This animal presents a specific morphology that may present challenges to this neural pathway in terms of the metabolism required for correct functioning and maintenance of potentially very long axons. The spinal cord of the giraffe can be up to 2.6 m long and forms the conus medullaris at the level of the sacral vertebrae. Primary motor cortex was found in a location typical of that of other ungulates, and the cytoarchitectonic appearance of this cortical area was similar to that previously reported for sheep, despite the potential distance that the axons emanating from the layer 5 gigantopyramidal neurons must travel. A typically mammalian dorsal striatopallidal complex was transected by a strongly coalesced internal capsule passing through to the pons and forming clearly identifiable but somewhat flattened (in a dorsoventral plane) pyramidal tracts. These tracts terminated in a spinal cord that exhibited no unique anatomical features related to its length. Our results, at least at the level of organization investigated herein, show that the corticospinal tract of the giraffe resembled that of a typical ungulate.  相似文献   

14.
Summary Postnatal development of the expression and localization of a membrane-associated cytoskeletal protein, calspectin (fodrin or brain spectrin), in the visual cortex, was immunohistochemically studied in newborn to adult rats, by using an anti-calspectin antibody. At birth, calspectin-immunoreactivity was already present at the plasma membrane and in the cytoplasm of neurons which were mostly pyramidal cells located in the upper part of the cortical subplate. Immature neurons located in the cortical plate were not stained by the antibody, suggesting that calspectin is expressed only in neurons which have differentiated or are differentiating.At postnatal days 2 to 7, immunoreactive neurons were dramatically increased in layers V and VI and very intense labelling was seen in the apical dendrites of layer V pyramidal cells. Most of the stained processes of these and other neurons showed signs of rapid dendritic growth, i.e. non-terminal as well as terminal growth cones and filopodia. At days 10 to 17, dendrites of pyramidal cells in layers II and III became clearly detectable, although still slender. At days 24 to 34, the basal dendrites of pyramidal cells in layers II, III and V became intensely immunoreactive and dendritic spines were visualized by the antibody. In the adult, however, the calspectin immunoreactivity became very weak and spines were not recognizable. At all the ages, axons and neuroglia were unstained. Also, most of the neurons in layer IV of the cortex were not immunoreactive.These results suggest that caispectin is most abundantly expressed in growing parts of the dendrites and spines. A hypothesis that calspectin may play a role in synaptic plasticity in the developing visual cortex is discussed.  相似文献   

15.
Summary The North American opossum is born 12 days after conception and is therefore available for experimental manipulation in an immature state. We have used the opossum to study the growth of cortical axons into the brainstem and spinal cord and have obtained evidence that such growth occurs in an orderly fashion. Cortical axons reach the ventral mesencephalon 12 days after birth and some of them have grown into the caudal medulla where they decussate by 23 days. At the latter stage immature cortical axons also distribute to the midbrain tegmentum, the basilar pons, the inferior olive and the hilum of the nucleus cuneatus. Cortical axons first enter the spinal cord about 30 days after birth where they are present in the white matter before growing into the dorsal horn. The forelimb placing reaction does not develop until well after cortical axons have reached cervical levels. Axons from the cerebral cortex grow into the spinal cord before there is evidence for cortical innervation of either the red nucleus or the bulbar reticular formation and well before pyramidal cells of the neocortex are mature. The relatively late development of corticospinal and corticobulbar systems contrasts markedly with the early growth of bulbospinal axons.This investigation was supported by NS-07410 and BNS-8008675 as well as the WVa Medical Corporation  相似文献   

16.
Collateral sprouting of undamaged descending axons is one potential mechanism for recovery of function after incomplete spinal cord injury. In this study, we have investigated whether terminals of the intact corticospinal tract in the rat would sprout following ablation of a parallel descending pathway, the rubrospinal tract. No sprouting was detected after this injury alone. However, the combination of rubrospinal tract ablation with administration of 100ng neurotrophin-3 to neurons of the corticospinal tract resulted in marked increased density of corticospinal innervation in the superficial dorsal horn. There was no effect of administration of neurotrophin-3 alone and increase in axon density was not detected in the deep dorsal horn.These results imply that spontaneous sprouting of undamaged corticospinal axons does not occur following ablation of a parallel tract system, although collateral sprouting can be induced through a combination of the lesion plus exogenous growth factor. Induced change in corticospinal terminal density is detected in the superficial dorsal horn only, supporting the hypothesis that this is an area particularly supportive of circuit reorganisation.  相似文献   

17.
The expression of the growth associated protein, GAP-43, in developing rat hindlimb peripheral nerves has been studied using immunocytochemistry. GAP-43, is first detected in lumbar spinal nerves at embryonic day (E)12 as the axons grow to the base of the hindlimb. It is expressed along the whole length of the nerves as well as in the growth cones. GAP-43 staining becomes very intense over the next 36 h while the axons remain in the plexus region at the base of the limb bud before forming peripheral nerves at E14. It remains intense along the length of the growing peripheral nerves, the first of which are cutaneous, branching away from the plexus and growing specifically to the skin, their axon tips penetrating the epidermis of the proximal skin at E15 and the toes at E19. GAP-43-containing terminals form a dense plexus throughout the epidermis which subsequently withdraws subepidermally in the postnatal period. GAP-43 staining is also evident along the growing muscle nerves during muscle innervation, which follows behind that of skin. Axons branch over the surface of proximal muscles at E15 but do not form terminals until E17. As target innervation proceeds, GAP-43 staining declines in the proximal part of the nerve but remains intense in the distal portions. Overall GAP-43 expression in the hindlimb decreases in the second postnatal week as axon growth and peripheral terminal formation decline.  相似文献   

18.
Reconstruction of dendritic growth cones in neonatal mouse olfactory bulb   总被引:3,自引:0,他引:3  
Summary An improved and simplified method of serial sectioning for electron microscopy was used to prepare material from which were reconstructed dendritic growth cones of mitral cells in neonatal mouse olfactory glomerulus. Growth cones are characterized by: (1) one or more filopodia, each approximately 0.2 m in diameter, projecting from an expanded dendritic terminal or pre-terminal region; (2) a polygonal array of microfilaments approximately 50 Å in diameter, filling the enlarged terminal and filopodia; (3) an absence of microtubules and a paucity of mitochondria; (4) a few profiles of smooth endoplasmic reticulum or vesicles; (5) occasional axodendritic synapses. These characteristics are compared with those of other growing processes in the nervous system and a consistent picture of the appearance of growing neuronal processes obtained, suitable to use as a guide in a search for additional growing processes in developing and mature brains.  相似文献   

19.
背景:有关皮质脊髓束受损后损伤附近及远处超微结构的整体变化的报道甚少。 目的:观察大鼠皮质脊髓束半横断损伤后皮质脊髓束超微结构的变化。 方法:选择性切割SD大鼠延髓左侧锥体,建立皮质脊髓束半横断损伤模型,于造模后4,14,28 d取受损皮质脊髓束,制备电镜标本。 结果与结论:透射电镜观察发现受损皮质脊髓束髓鞘和轴索发生肿胀,形态不规则。随着时间的延长,溃变进行性加重,受损皮质脊髓束主要表现为髓鞘破坏、溶解及轴索脱髓鞘病变、胞浆浓缩、细胞器增多及空泡样变性。脊髓颈、胸、腰段受损皮质脊髓束的溃变比损伤部位严重。  相似文献   

20.
Intra- or juxta-columnar connections of pyramidal neurons to corticospinal neurons in rat motorsensory cortices were examined with brain slices by combining intracellular staining with Golgi-like retrograde labeling of corticospinal neurons. Of 108 intracellularly labeled pyramidal neurons, 27 neurons were selected for morphological analysis by successful staining of their axonal arborizations and sufficient retrograde labeling of corticospinal neurons. Many varicosities of local axon collaterals of each pyramidal neuron were closely apposed to the dendrites of corticospinal neurons, suggesting the convergent projections of layer II–VI pyramidal neurons to corticospinal neurons. Particularly, the varicosities of a layer IV star-pyramidal neuron made two- to three-fold more appositions to the dendrites of corticospinal neurons than those of a pyramidal neuron in the other layers. Fifteen appositions were examined electron-microscopically and 60% of them made asymmetric axospinous synapses. The present results together with those of the preceding report suggest that thalamic inputs are conveyed to corticospinal neurons preferentially via layer IV star-pyramidal neurons with phasic response properties, and thereby might contribute to the initiation or switching of movement. In contrast, inputs with tonic response properties from the other layers seem to be integrated in corticospinal neurons, and might be useful in maintaining the activity of corticospinal neurons.  相似文献   

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