Hawkins R D, Lalevic N, Clark G A, Kandel E R
Center for Neurobiology and Behavior, College of Physicians and Surgeons, Columbia University, New York, NY.
Proc Natl Acad Sci U S A. 1989 Oct;86(19):7620-4. doi: 10.1073/pnas.86.19.7620.
The gill- and siphon-withdrawal reflex of Aplysia undergoes classical conditioning of its amplitude and duration when siphon stimulation (the conditioned stimulus, CS) is paired with tail or mantle shock (the unconditioned stimulus, US). This conditioning of a preexisting response exhibits both temporal and stimulus specificities, which can be accounted for by activity-dependent enhancement of presynaptic facilitation of the siphon sensory neurons. To test whether conditioning of the reflex also exhibits response specificity (development of a new type of response to the CS that often resembles the response to the US), we measured the direction of siphon withdrawal in response to siphon stimulation (the CS) with tail or mantle shock as the US. The unlearned response to siphon stimulation is straight contraction, the response to tail shock is backward bending, and the response to mantle shock is forward bending. In the first experiment, we trained different animals with the tail or mantle US paired or unpaired with the CS; in a second experiment, we trained each animal with two CSs, one of which was paired with the US; in a third experiment, we varied US intensity; and in a fourth experiment, we trained each animal with two USs, one of which was paired with the CS. There was a significant, pairing-specific tendency for the direction of the response to the CS to resemble the response to the US after training in each experiment, demonstrating response specificity in conditioning of the withdrawal reflex. This feature of conditioning could in principle be accounted for by an elaboration of activity-dependent facilitation.
当虹吸管刺激(条件刺激,CS)与尾部或外套膜电击(非条件刺激,US)配对时,海兔的鳃和虹吸管退缩反射在其幅度和持续时间上会经历经典条件作用。这种对预先存在反应的条件作用表现出时间和刺激特异性,这可以通过虹吸管感觉神经元突触前易化的活动依赖性增强来解释。为了测试反射的条件作用是否也表现出反应特异性(对CS产生一种通常类似于对US反应的新型反应),我们以尾部或外套膜电击作为US,测量了对虹吸管刺激(CS)的虹吸管退缩方向。对虹吸管刺激的未学习反应是直接收缩,对尾部电击的反应是向后弯曲,对外套膜电击的反应是向前弯曲。在第一个实验中,我们用与CS配对或不配对的尾部或外套膜US训练不同的动物;在第二个实验中,我们用两个CS训练每只动物,其中一个与US配对;在第三个实验中,我们改变US强度;在第四个实验中,我们用两个US训练每只动物,其中一个与CS配对。在每个实验中训练后,对CS的反应方向都有显著的、配对特异性的倾向,类似于对US的反应,这表明在退缩反射的条件作用中存在反应特异性。原则上,这种条件作用的特征可以通过对活动依赖性易化的细化来解释。