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TaggedAPTARAFigure TaggedAPTARAEndEffect of exercise on locomotor system                              161






































           Fig. 1. Exercise-induced adaptation of neurons in the locomotor system. The treadmill exercise affects neurons in the CPG and MLR. The plasticity of neurons
           includes (A) modulated ion channels, (B) promoted dendritic plasticity, (C) increased number of excitatory synapses, and (D) increased neurotrophic expression.
           CPG = central pattern generator; DRG = dorsal root ganglion; MLR = mesencephalic locomotor region.TaggedAPTARAEnd
           potential (RMP), input resistance (Rin, calculated by  motoneurons. 8,1820  Specifically, their lumbar motoneurons
           membrane potential deflection divided by negatively injected  undergo a reduction in Rin 20,21  followed by a decrease in ampli-
           current), frequency and current relationship (FI relationship,  tude of AHP during similar interspike intervals. 22  Additional
           established by firing frequency vs. step current), action poten-  studies of fictive locomotor outputs in cat motoneurons have also
           tial (AP), and afterhyperpolarization (AHP, hyperpolarization  revealed significant changes in their FI relationship, a pattern
           of membrane potential after an AP). The ion channels focused  of nonlinear voltage-dependent excitation, and perhaps most
           on in this review included: (a) transient sodium channels,  significantly, a hyperpolarization of their Vth required for AP
           which determine the threshold of AP generation, (b) persistent  generation. 8,9,2227  The varying changes in FI relationship
           sodium channels, which generate persistent inward currents  during fictive locomotion are shown as the left-shift of FI rela-
           (PICs) important for motoneuron excitability, (c) L-type calcium  tionship with alteration of FI slopes during the excitatory phase
           channels, which generate PICs regulating rhythm generation in  of the locomotor drive potentials and the right-shift of FI rela-
           the locomotor system, (d) delayed-rectifier potassium (K(DR))  tionship with changes in FI slopes in the inhibitory phase of the
           channels, which modulate Vth, and (e) calcium-activated potas-  locomotor drive potentials. In addition, the disappearance of FI
                                                                 relationships in the excitatory phase of the locomotor drive poten-
           sium channels (K(Ca)), which mediate AHP.TaggedAPTARAEnd
                                                                 tials is also observed in some cells, where the motoneuron
           TaggedAPTARAH12. Acute exercise increases neuronal excitabilityTaggedAPTARAEnd  discharge rates are saturated without changing with the injected
             TaggedAPTARAPOur present understanding of the relationship between acute  currents. 22,24  Similar alterations in neuronal membrane properties
           exercise and neuronal excitability has emerged from comprehen-  indicating acute exercise-enhanced neuronal excitability have
           sive studies of fictive locomotion in different animal models.  also been reported in other species. Studies of fictive locomotion
           Fictive locomotion refers to the generation of neural activity  in decerebrated neonatal rats likewise demonstrate that their
           patterns in the nervous system resembling those patterns  spinal motoneurons undergo decreases in amplitude of AHP and
           observed during actual locomotion, but without actual movement  a hyperpolarization of their Vth during locomotor-like
                                                                       9,26,28
           of the limbs or body; hence, the resulting locomotion is consid-  activity.  Increases in neuronal excitability experienced
           ered not real but fictive. 17  Studies of fictive locomotion in cats  during acute exercise ultimately dissipate back into a steady state
           have demonstrated that acute exercise leads to enhanced neuronal  once the locomotor state terminates such that the cycle may
           excitability. For instance, electrical stimulation of mesencephalic  repeat again. Mechanism studies suggest that ion channels are
           locomotor regions of decerebrated cats generates locomotion 17  responsible for regulating this cycle. Experimental and modeling
           via  the  increase  of  the  excitability  of  their  spinal  studies indicate that the hyperpolarization of Vth can be mediated
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