e., laminar) fMRI could potentially be used to address such questions. The six cortical layers have different distributions of cell types, cell sizes, connectivity, energy use, etc., reflecting the different functions of the layers. For instance, Bleomycin supplier input to a cortical area typically arrives in layer IV, while output is typically generated in layer V. In primary visual

cortex (V1), the different stimulus selectivity of the layers, e.g., the magno- and parvocellular pathways, is well known (Callaway, 1998). The relative thickness of the layers also varies for different cortical areas depending on the function of the area. If these anatomical and functional differences have a counterpart in the fMRI signals, fMRI at laminar resolution might be used to elucidate such different cortical computations. However, laminar differences under different stimulus conditions have remained elusive. There could be multiple reasons for this, for instance resolution limitations. Another possible reason is that the profile of the BOLD response as a function of cortical depth could be determined by the properties of the vasculature, with the laminar profile

of the BOLD response only exhibiting amplitude differences, independent of which layers show strongest neural activity. Yet another possibility is that the point spread function (PSF) of the hemodynamic Everolimus chemical structure response is larger than the thickness of the layers. To address these questions and to investigate whether obvious laminar differences in the patterns of the BOLD response exist, we compared the laminar properties of positive and negative BOLD responses. We chose these stimuli because of the large differences between the responses and because the negative BOLD signal has been reported to have different properties; namely, to be more specific than the positive BOLD signal (Bressler et al., 2007) and to provide independent information about brain function (Wade and Rowland, 2010). Negative

BOLD responses have been observed in humans and animals (Allison et al., 2000; Harel et al., 2002; Huang et al., 1996; Shmuel et al., 2002, 2006; Resminostat Tootell et al., 1998). In V1, negative responses can be reliably observed adjacent to positive BOLD signals (Huang et al., 1996; Shmuel et al., 2002, 2006; Tootell et al., 1998; Wade and Rowland, 2010). They were also observed upon ipsilateral inhibition in visual-, motor- and somatosensory cortex (Allison et al., 2000; Hlushchuk and Hari, 2006; Schäfer et al., 2012; Smith et al., 2004; Stefanovic et al., 2004; Whittingstall et al., 2008). Negative BOLD signals were shown to be associated with decreases in cerebral blood flow (CBF) and neural activity (Boorman et al., 2010; Devor et al., 2007; Shmuel et al., 2002, 2006).