Bourgeois J P, Popot J L, Ryter A, Changeux J P
J Cell Biol. 1978 Oct;79(1):200-16. doi: 10.1083/jcb.79.1.200.
Electroplaques dissected from the electric organ of Electrophorus electricus are labeled by tritiated alpha1-isotoxin from Naja nigricollis, a highly selective reagent of the cholinergic (nicotinic) receptor site. Preincubation of the cell with an excess of unlabeled alpha-toxin and with a covalent affinity reagent or labeling in the presence of 10(-4) M decamethonium reduces the binding of [3H]alpha-toxin by at least 75%. Absolute surface densities of alpha-toxin sites are estimated by high-resolution autoradiography on the basis of silver grain distribution and taking into account the complex geopmetry of the cell surface. Binding of [3H]alpha-toxin on the noninnervated face does not differ from background. Labeled sites are observed on the innervated membrane both between the synapses and under the nerve terminals but the density of sites is approx. 100 times higher at the level of the synapses than in between. Analysis of the distance of silver grains from the innervated membrane shows a symmetrical distribution centered on the postsynaptic plasma membrane under the nerve terminal. In extrasynaptic areas, the barycenter of the distribution lies approximately 0.5 micrometer inside the cell, indicating that alpha-toxin sites are present on the membrane of microinvaginations, or caveolae, abundant in the extrajunctional areas. An absolute density of 49,600 +/- 16,000 sites/micrometer2 of postsynaptic membrane is calculated; it is in the range of that found at the crest of the folds at the neuromuscular junction and expected from a close packing of receptor molecules. Electric organs were denervated for periods up to 142 days. Nerve transmission fails after 2 days, and within a week all the nerve terminals disappear and are subsequently replaced by Schwann cell processes, whereas the morphology of the electroplaque remains unaffected. The denervated electroplaque develops some of the electrophysiological changes found with denervated muscles (increases of membrane resting resistance, decrease of electrical excitability) but does not become hypersensitive to cholinergic agonists. Autoradiography of electroplaques dissected from denervated electric organs reveals, after labeling with [3H]alpha-toxin, patches of silver grains with a surface density close to that found in the normal electroplaque. The density of alpha-toxin binding sites in extrasynaptic areas remains close to that observed on innervated cells, confirming that denervation does not cause an increase in the number of cholinergic receptor sites. The patches have the same distribution, shape,and dimensions as in subneural areas of the normal electroplaque, and remnants of nerve terminal or Schwann cells are often found at the level of the patches. They most likely correspond to subsynaptic areas which persist with the same density of [3H]alpha-toxin sites up to 52 days after denervation. In the adult synapse, therefore, the receptor protein exhibits little if any tendency for lateral diffusion.
从电鳗(Electrophorus electricus)的电器官中分离出的电板,可被来自黑颈眼镜蛇(Naja nigricollis)的氚化α1-毒素标记,该毒素是胆碱能(烟碱型)受体位点的高度选择性试剂。用过量的未标记α-毒素和共价亲和试剂对细胞进行预孵育,或在10⁻⁴ M十烃季铵存在下进行标记,可使[³H]α-毒素的结合减少至少75%。基于银颗粒分布并考虑到细胞表面复杂的几何形状,通过高分辨率放射自显影术估计α-毒素位点的绝对表面密度。[³H]α-毒素在非神经支配面的结合与背景无差异。在神经支配膜上,突触之间和神经末梢下方均观察到标记位点,但突触处的位点密度比其间约高100倍。对银颗粒与神经支配膜距离的分析显示,以神经末梢下方的突触后质膜为中心呈对称分布。在突触外区域,分布的重心位于细胞内约0.5微米处,表明α-毒素位点存在于突触外区域丰富的微内陷或小窝的膜上。计算出突触后膜的绝对密度为49,600±16,000个位点/微米²;这在神经肌肉接头褶皱嵴处发现的范围内,并且符合受体分子紧密堆积的预期。将电器官去神经支配长达142天。2天后神经传递失败,一周内所有神经末梢消失,随后被施万细胞突起取代,而电板的形态不受影响。去神经支配的电板出现了一些去神经支配肌肉所具有的电生理变化(膜静息电阻增加,电兴奋性降低),但对胆碱能激动剂不变得超敏。用[³H]α-毒素标记后,对从去神经支配的电器官中分离出的电板进行放射自显影,发现银颗粒斑块的表面密度与正常电板中发现的接近。突触外区域α-毒素结合位点的密度仍与神经支配细胞上观察到的接近,证实去神经支配不会导致胆碱能受体位点数量增加。这些斑块与正常电板神经下区域具有相同的分布、形状和尺寸,并且在斑块水平经常发现神经末梢或施万细胞的残余物。它们很可能对应于突触下区域,在去神经支配后长达52天内,这些区域的[³H]α-毒素位点密度保持不变。因此,在成年突触中,受体蛋白几乎没有横向扩散的趋势。