Brembs Björn
Freie Universität Berlin, Institute for Biology-Neurobiology, Berlin, Germany.
Behav Processes. 2011 May;87(1):157-64. doi: 10.1016/j.beproc.2011.02.005. Epub 2011 Mar 15.
Already in the 1930s Skinner, Konorski and colleagues debated the commonalities, differences and interactions among the processes underlying what was then known as "conditioned reflexes type I and II", but which is today more well-known as classical (Pavlovian) and operant (instrumental) conditioning. Subsequent decades of research have confirmed that the interactions between the various learning systems engaged during operant conditioning are complex and difficult to disentangle. Today, modern neurobiological tools allow us to dissect the biological processes underlying operant conditioning and study their interactions. These processes include initiating spontaneous behavioral variability, world-learning and self-learning. The data suggest that behavioral variability is generated actively by the brain, rather than as a by-product of a complex, noisy input-output system. The function of this variability, in part, is to detect how the environment responds to such actions. World-learning denotes the biological process by which value is assigned to environmental stimuli. Self-learning is the biological process which assigns value to a specific action or movement. In an operant learning situation using visual stimuli for flies, world-learning inhibits self-learning via a prominent neuropil region, the mushroom-bodies. Only extended training can overcome this inhibition and lead to habit formation by engaging the self-learning mechanism. Self-learning transforms spontaneous, flexible actions into stereotyped, habitual responses.
早在20世纪30年代,斯金纳、科诺尔斯基及其同事就对当时被称为“第一型和第二型条件反射”(如今更熟知的是经典(巴甫洛夫式)和操作性(工具性)条件作用)背后的过程的共性、差异和相互作用展开了辩论。随后几十年的研究证实,在操作性条件作用过程中涉及的各种学习系统之间的相互作用是复杂且难以厘清的。如今,现代神经生物学工具使我们能够剖析操作性条件作用背后的生物学过程并研究它们的相互作用。这些过程包括引发自发行为变异性、环境学习和自我学习。数据表明,行为变异性是由大脑主动产生的,而非复杂、嘈杂的输入输出系统的副产品。这种变异性的功能部分在于检测环境对这类行为的反应。环境学习指的是给环境刺激赋予价值的生物学过程。自我学习是给特定行为或动作赋予价值的生物学过程。在利用视觉刺激对果蝇进行的操作性学习情境中,环境学习通过一个突出的神经纤维区域——蘑菇体抑制自我学习。只有经过长时间训练才能克服这种抑制,并通过激活自我学习机制导致习惯形成。自我学习将自发、灵活的行为转变为刻板的习惯性反应。