Kucera J, Walro J M, Reichler J
Department of Neurology, School of Medicine, Boston University, Massachusetts 02118.
Am J Anat. 1988 Dec;183(4):344-58. doi: 10.1002/aja.1001830408.
The chronology of development of spindle neural elements was examined by electron microscopy in fetal and neonatal rats. The three types of intrafusal muscle fiber of spindles from the soleus muscle acquired sensory and motor innervation in the same sequence as they formed--bag2, bag1, and chain. Both the primary and secondary afferents contacted developing spindles before day 20 of gestation. Sensory endings were present on myoblasts, myotubes, and myofibers in all intrafusal bundles regardless of age. The basic features of the sensory innervation--first-order branching of the parent axon, separation of the primary and secondary sensory regions, and location of both primary and secondary endings beneath the basal lamina of the intrafusal fibers--were all established by the fourth postnatal day. Cross-terminals, sensory terminals shared by more than one intrafusal fiber, were more numerous at all developmental stages than in mature spindles. No afferents to immature spindles were supernumerary, and no sensory axons appeared to retract from terminations on intrafusal fibers. The earliest motor axons contacted spindles on the 20th day of gestation or shortly afterward. More motor axons supplied the immature spindles, and a greater number of axon terminals were visible at immature intrafusal motor endings than in adult spindles; hence, retraction of supernumerary motor axons accompanies maturation of the fusimotor system analogous to that observed during the maturation of the skeletomotor system. Motor endings were observed only on the relatively mature myofibers; intrafusal myoblasts and myotubes lacked motor innervation in all age groups. This independence of the early stages of intrafusal fiber assembly from motor innervation may reflect a special inherent myogenic potential of intrafusal myotubes or may stem from the innervation of spindles by sensory axons.
通过电子显微镜检查了胎鼠和新生大鼠中梭形神经元件的发育时间顺序。比目鱼肌梭内的三种类型的肌梭内肌纤维在形成时按照相同的顺序获得感觉和运动神经支配——袋状2型、袋状1型和链状。初级和次级传入纤维在妊娠第20天之前就接触到了发育中的肌梭。无论年龄大小,所有肌梭内束的成肌细胞、肌管和肌纤维上都有感觉末梢。感觉神经支配的基本特征——母轴突的一级分支、初级和次级感觉区域的分离以及初级和次级末梢在肌梭内纤维基膜下方的位置——在出生后第4天就已全部形成。交叉末梢,即由不止一根肌梭内纤维共享的感觉末梢,在所有发育阶段都比成熟肌梭中的更多。未成熟肌梭没有多余的传入纤维,也没有感觉轴突似乎从肌梭内纤维的终末缩回。最早的运动轴突在妊娠第20天或之后不久接触到肌梭。更多的运动轴突支配未成熟的肌梭,并且在未成熟的肌梭内运动终末可见的轴突终末数量比成年肌梭中的更多;因此,多余运动轴突的缩回伴随着肌梭运动系统的成熟,这与在骨骼运动系统成熟过程中观察到的情况类似。运动终末仅在相对成熟度的肌纤维上观察到;所有年龄组的肌梭内成肌细胞和肌管都缺乏运动神经支配。肌梭内纤维组装早期阶段与运动神经支配的这种独立性可能反映了肌梭内肌管特殊的内在肌源性潜力,或者可能源于感觉轴突对肌梭的神经支配。