McGill University, Montreal Neurological Institute, 3801 University St., Montreal, QC H3A 2B4, Canada.
J Neurophysiol. 2010 Jun;103(6):3123-38. doi: 10.1152/jn.00654.2009. Epub 2010 Mar 24.
Neurons in the primate extrastriate cortex are highly selective for complex stimulus features such as faces, objects, and motion patterns. One explanation for this selectivity is that neurons in these areas carry out sophisticated computations on the outputs of lower-level areas such as primary visual cortex (V1), where neuronal selectivity is often modeled in terms of linear spatiotemporal filters. However, it has long been known that such simple V1 models are incomplete because they fail to capture important nonlinearities that can substantially alter neuronal selectivity for specific stimulus features. Thus a key step in understanding the function of higher cortical areas is the development of realistic models of their V1 inputs. We have addressed this issue by constructing a computational model of the V1 neurons that provide the strongest input to extrastriate cortical middle temporal (MT) area. We find that a modest elaboration to the standard model of V1 direction selectivity generates model neurons with strong end-stopping, a property that is also found in the V1 layers that provide input to MT. With this computational feature in place, the seemingly complex properties of MT neurons can be simulated by assuming that they perform a simple nonlinear summation of their inputs. The resulting model, which has a very small number of free parameters, can simulate many of the diverse properties of MT neurons. In particular, we simulate the invariance of MT tuning curves to the orientation and length of tilted bar stimuli, as well as the accompanying temporal dynamics. We also show how this property relates to the continuum from component to pattern selectivity observed when MT neurons are tested with plaids. Finally, we confirm several key predictions of the model by recording from MT neurons in the alert macaque monkey. Overall our results demonstrate that many of the seemingly complex computations carried out by high-level cortical neurons can in principle be understood by examining the properties of their inputs.
灵长类动物外纹状皮层中的神经元对复杂刺激特征(如人脸、物体和运动模式)具有高度选择性。这种选择性的一种解释是,这些区域中的神经元对初级视觉皮层 (V1) 等低级区域的输出进行了复杂的计算,而神经元的选择性通常以线性时空滤波器来建模。然而,长期以来人们一直知道,这种简单的 V1 模型是不完整的,因为它们未能捕捉到重要的非线性,这些非线性可以极大地改变神经元对特定刺激特征的选择性。因此,理解高级皮层区域功能的关键步骤是开发其 V1 输入的现实模型。我们通过构建一个为外纹状皮层中的中颞(MT)区提供最强输入的 V1 神经元的计算模型来解决这个问题。我们发现,对 V1 方向选择性的标准模型进行适度的改进,可以生成具有强烈端抑制特性的模型神经元,这种特性也存在于为 MT 提供输入的 V1 层中。有了这个计算特征,只需假设它们对输入进行简单的非线性求和,就可以模拟 MT 神经元看似复杂的特性。这个模型具有非常少的自由参数,可以模拟 MT 神经元的许多不同特性。特别是,我们模拟了 MT 调谐曲线对倾斜棒刺激的方向和长度的不变性,以及伴随的时间动态。我们还展示了这种特性如何与当 MT 神经元用光栅进行测试时观察到的从成分选择性到模式选择性的连续体相关。最后,我们通过在警觉的猕猴中记录 MT 神经元来验证模型的几个关键预测。总的来说,我们的结果表明,许多高级皮层神经元进行的看似复杂的计算原则上可以通过检查其输入的特性来理解。