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A quantitative computational theory of the operation of the hippocampus as an episodic memory system is described. The CA3 system operates as a single attractor or autoassociation network (1) to enable rapid one-trial associations between any spatial location (place in rodents or spatial view in primates) and an object or reward and (2) to provide for completion of the whole memory during recall from any part. The theory is extended to associations between time and object or reward to implement temporal order memory, which is also important in episodic memory. The dentate gyrus performs pattern separation by competitive learning to create sparse representations producing, for example, neurons with place-like fields from entorhinal cortex grid cells. The dentate granule cells generate, by the very small number of mossy fibre connections to CA3, a randomizing pattern separation effect that is important during learning but not recall and that separates out the patterns represented by CA3 firing as being very different from each other. This is optimal for an unstructured episodic memory system in which each memory must be kept distinct from other memories. The direct perforant path input to CA3 is quantitatively appropriate for providing the cue for recall in CA3 but not for learning. The CA1 recodes information from CA3 to set up associatively learned backprojections to the neocortex to allow the subsequent retrieval of information to the neocortex, giving a quantitative account of the large number of hippocampo-neocortical and neocortical-neocortical backprojections. Tests of the theory including hippocampal subregion analyses and hippocampal NMDA receptor knockouts are described and support the theory.

An alternative parcellation of the orbitofrontal cortex is described for the automated anatomical labeling atlas of Tzourio-Mazoyer et al. (2002) (Automated anatomical labeling of activations in SPM using a macroscopic anatomical parcellation of the MNI MRI single-subject brain. NeuroImage 15:273–289). The new parcellation of the orbitofrontal cortex follows the description provided by Chiavaras, Petrides, and colleagues (2000, 2001). The new atlas is available as a toolbox for SPM at http://www.gin.cnrs.fr/AAL2. •A new parcellation of the orbitofrontal cortex for the AAL•The AAL is the automated anatomical labeling atlas of Tzourio-Mazoyer et al. (2002).•The new atlas, AAL2, is available as aal2.nii.gz at http://www.gin.cnrs.fr.

Following a first version AAL of the automated anatomical labeling atlas (Tzourio-Mazoyer et al., 2002), a second version (AAL2) (Rolls et al., 2015) was developed that provided an alternative parcellation of the orbitofrontal cortex following the description provided by Chiavaras, Petrides, and colleagues. We now provide a third version, AAL3, which adds a number of brain areas not previously defined, but of interest in many neuroimaging investigations. The 26 new areas in the third version are subdivision of the anterior cingulate cortex into subgenual, pregenual and supracallosal parts; subdivision of the thalamus into 15 parts; the nucleus accumbens, substantia nigra, ventral tegmental area, red nucleus, locus coeruleus, and raphe nuclei. The new atlas is available as a toolbox for SPM, and can be used with MRIcron. •The automated anatomical atlas 3 (AAL3) is described. The following new areas are added.•Subdivision of the anterior cingulate cortex into subgenual, pregenual and supracallosal parts.•Thalamus, nucleus accumbens, substantia nigra, ventral tegmental area, red nucleus.•Locus coeruleus, and raphe nuclei.•AAL3 is available as a toolbox for SPM at www.oxcns.org.

The human posterior cingulate, retrosplenial, and medial parietal cortex are involved in memory and navigation. The functional anatomy underlying these cognitive functions was investigated by measuring the effective connectivity of these Posterior Cingulate Division (PCD) regions in the Human Connectome Project‐MMP1 atlas in 171 HCP participants, and complemented with functional connectivity and diffusion tractography. First, the postero‐ventral parts of the PCD (31pd, 31pv, 7m, d23ab, and v23ab) have effective connectivity with the temporal pole, inferior temporal visual cortex, cortex in the superior temporal sulcus implicated in auditory and semantic processing, with the reward‐related vmPFC and pregenual anterior cingulate cortex, with the inferior parietal cortex, and with the hippocampal system. This connectivity implicates it in hippocampal episodic memory, providing routes for “what,” reward and semantic schema‐related information to access the hippocampus. Second, the antero‐dorsal parts of the PCD (especially 31a and 23d, PCV, and also RSC) have connectivity with early visual cortical areas including those that represent spatial scenes, with the superior parietal cortex, with the pregenual anterior cingulate cortex, and with the hippocampal system. This connectivity implicates it in the “where” component for hippocampal episodic memory and for spatial navigation. The dorsal–transitional–visual (DVT) and ProStriate regions where the retrosplenial scene area is located have connectivity from early visual cortical areas to the parahippocampal scene area, providing a ventromedial route for spatial scene information to reach the hippocampus. These connectivities provide important routes for “what,” reward, and “where” scene‐related information for human hippocampal episodic memory and navigation. The midcingulate cortex provides a route from the anterior dorsal parts of the PCD and the supracallosal part of the anterior cingulate cortex to premotor regions. First, the postero‐ventral parts of the Posterior Cingulate Division of the Human Connectome Project Multimodal Parcellation atlas (PCD; 31pd, 31pv, 7m, d23ab, and v23ab) have effective connectivity with the temporal pole, inferior temporal visual cortex, cortex in the superior temporal sulcus implicated in auditory and semantic processing, with the reward‐related vmPFC and pregenual anterior cingulate cortex, with the inferior parietal cortex, and with the hippocampal system; and this connectivity implicates these regions in hippocampal episodic memory, providing routes for “what,” reward, and semantic schema‐related information to access the hippocampus. Second, the antero‐dorsal parts of the PCD (especially 31a and 23d, PCV, and also RSC) have connectivity with early visual cortical areas including those that represent spatial scenes, with the superior parietal cortex, with the pregenual anterior cingulate cortex, and with the hippocampal system. This connectivity implicates them in the “where” component for hippocampal episodic memory and for spatial navigation. Third, the dorsal–transitional–visual (DVT) and ProStriate regions are where the retrosplenial scene area is located, have connectivity from early cortical visual areas, connect to the parahippocampal scene area, and provide a ventromedial cortex route for scene information to reach the hippocampal system.

The local recurrent collateral connections between cortical neurons provide a basis for attractor neural networks for memory, attention, decision-making, and thereby for many aspects of human behavior. In schizophrenia, a reduction of the firing rates of cortical neurons, caused for example by reduced NMDA receptor function or reduced spines on neurons, can lead to instability of the high firing rate attractor states that normally implement short-term memory and attention in the prefrontal cortex, contributing to the cognitive symptoms. Reduced NMDA receptor function in the orbitofrontal cortex by reducing firing rates may produce negative symptoms, by reducing reward, motivation, and emotion. Reduced functional connectivity between some brain regions increases the temporal variability of the functional connectivity, contributing to the reduced stability and more loosely associative thoughts. Further, the forward projections have decreased functional connectivity relative to the back projections in schizophrenia, and this may reduce the effects of external bottom-up inputs from the world relative to internal top-down thought processes. Reduced cortical inhibition, caused by a reduction of GABA neurotransmission, can lead to instability of the spontaneous firing states of cortical networks, leading to a noise-induced jump to a high firing rate attractor state even in the absence of external inputs, contributing to the positive symptoms of schizophrenia. In depression, the lateral orbitofrontal cortex non-reward attractor network system is over-connected and has increased sensitivity to non-reward, providing a new approach to understanding depression. This is complemented by under-sensitivity and under-connectedness of the medial orbitofrontal cortex reward system in depression.

To advance understanding of brain networks involved in language, the effective connectivity between 26 cortical regions implicated in language by a community analysis and 360 cortical regions was measured in 171 humans from the Human Connectome Project, and complemented with functional connectivity and diffusion tractography, all using the HCP multimodal parcellation atlas. A (semantic) network (Group 1) involving inferior cortical regions of the superior temporal sulcus cortex (STS) with the adjacent inferior temporal visual cortex TE1a and temporal pole TG, and the connected parietal PGi region, has effective connectivity with inferior temporal visual cortex (TE) regions; with parietal PFm which also has visual connectivity; with posterior cingulate cortex memory-related regions; with the frontal pole, orbitofrontal cortex, and medial prefrontal cortex; with the dorsolateral prefrontal cortex; and with 44 and 45 for output regions. It is proposed that this system can build in its temporal lobe (STS and TG) and parietal parts (PGi and PGs) semantic representations of objects incorporating especially their visual and reward properties. Another (semantic) network (Group 3) involving superior regions of the superior temporal sulcus cortex and more superior temporal lobe regions including STGa, auditory A5, TPOJ1, the STV and the Peri-Sylvian Language area (PSL) has effective connectivity with auditory areas (A1, A4, A5, Pbelt); with relatively early visual areas involved in motion, e.g., MT and MST, and faces/words (FFC); with somatosensory regions (frontal opercular FOP, insula and parietal PF); with other TPOJ regions; and with the inferior frontal gyrus regions (IFJa and IFSp). It is proposed that this system builds semantic representations specialising in auditory and related facial motion information useful in theory of mind and somatosensory / body image information, with outputs directed not only to regions 44 and 45, but also to premotor 55b and midcingulate premotor cortex. Both semantic networks (Groups 1 and 3) have access to the hippocampal episodic memory system via parahippocampal TF. A third largely frontal network (Group 2) (44, 45, 47l; 55b; the Superior Frontal Language region SFL; and including temporal pole TGv) receives effective connectivity from the two semantic systems, and is implicated in syntax and speech output.

A modified and extended version, HCPex, is provided of the surface-based Human Connectome Project-MultiModal Parcellation atlas of human cortical areas (HCP-MMP v1.0, Glasser et al. 2016). The original atlas with 360 cortical areas has been modified in HCPex for ease of use with volumetric neuroimaging software, such as SPM, FSL, and MRIcroGL. HCPex is also an extended version of the original atlas in which 66 subcortical areas (33 in each hemisphere) have been added, including the amygdala, thalamus, putamen, caudate nucleus, nucleus accumbens, globus pallidus, mammillary bodies, septal nuclei and nucleus basalis. HCPex makes available the excellent parcellation of cortical areas in HCP-MMP v1.0 to users of volumetric software, such as SPM and FSL, as well as adding some subcortical regions, and providing labelled coronal views of the human brain.