Vinay L, Brocard F, Pflieger J F, Simeoni-Alias J, Clarac F
CNRS, Développement et Pathologie du Mouvement, Marseille, France.
Brain Res Bull. 2000 Nov 15;53(5):635-47. doi: 10.1016/s0361-9230(00)00397-x.
The rat is quite immature at birth and a rapid maturation of motor behavior takes place during the first 2 postnatal weeks. Lumbar motoneurons undergo a rapid development during this period. The last week before birth represents the initial stages of motoneuron differentiation, including regulation of the number of cells and the arrival of segmental and first supraspinal afferents. At birth, motoneurons are electrically coupled and receive both appropriate and inappropriate connections from the periphery; the control from supraspinal structures is weak and exerted mainly through polysynaptic connections. During the 1st postnatal week, inappropriate sensori-motor contacts and electrical coupling disappear, the supraspinal control increases gradually and myelin formation is responsible for an increased conduction velocity in both descending and motor axons. Both N-methyl-D-aspartate (NMDA) and non-NMDA receptors are transiently overexpressed in the neonatal spinal cord. The contribution of non-NMDA receptors to excitatory amino acid transmission increases with age. Activation of gamma-aminobutyric acid(A) and glycine receptors leads to membrane depolarization in embryonic motoneurons but to hyperpolarization in older motoneurons. The firing properties of motoneurons change with development: they are capable of more repetitive firing at the end of the 1st postnatal week than before birth. However, maturation does not proceed simultaneously in the motor pools innervating antagonistic muscles; for instance, the development of repetitive firing of ankle extensor motoneurons lags behind that of flexor motoneurons. The spontaneous embryonic and neonatal network-driven activity, detected at the levels of motoneurons and primary afferent terminals, may play a role in neuronal maturation and in the formation and refinement of sensorimotor connections.
大鼠出生时相当不成熟,出生后的前两周运动行为会迅速成熟。在此期间,腰段运动神经元会经历快速发育。出生前的最后一周代表运动神经元分化的初始阶段,包括细胞数量的调节以及节段性和第一上位脊髓传入神经的到达。出生时,运动神经元通过电耦合连接,并且从外周接收合适和不合适的连接;上位脊髓结构的控制较弱,主要通过多突触连接发挥作用。在出生后的第一周,不合适的感觉运动接触和电耦合消失,上位脊髓控制逐渐增强,髓鞘形成导致下行轴突和运动轴突的传导速度增加。N-甲基-D-天冬氨酸(NMDA)受体和非NMDA受体在新生脊髓中均短暂过度表达。非NMDA受体对兴奋性氨基酸传递的贡献随年龄增加。γ-氨基丁酸A(GABA)受体和甘氨酸受体的激活在胚胎运动神经元中导致膜去极化,但在较成熟的运动神经元中导致超极化。运动神经元的放电特性随发育而变化:在出生后的第一周结束时,它们比出生前能够进行更多次重复放电。然而,支配拮抗肌的运动神经元池的成熟并非同时进行;例如,踝关节伸肌运动神经元重复放电的发育落后于屈肌运动神经元。在运动神经元和初级传入神经末梢水平检测到的自发胚胎和新生网络驱动活动,可能在神经元成熟以及感觉运动连接的形成和完善中发挥作用。