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D. Phil., School of Anatomical Sciences, Faculty of Health Sciences, University of the Witwatersrand, 2011 / Whether the neuropil is a static, optimally wired entity, whose components must be balanced in a certain way, is an open question. Are the proportions of the components of the neuropil consistent across different mammalian cortices, especially in primates where the cerebral cortex is complexly organized? This question is interesting because the actual biological underpinnings of complex behaviours and intelligence in big-brained primates remain enigmatic and why they seem qualitatively different from other animals in terms of their cognitive abilities. Understanding changes that may have occurred in the brain, especially at the level of neuropil organization, during the evolution in primates is important to our growing understanding of the intellectual abilities and behaviours exhibited by members of this group. The current series of quantitative studies was aimed at investigating variations in the proportionality of the “wire fraction” in three primate species, the olive baboon (Papio anubis), vervet monkey (Cercopithecus aethiops) and the common chimpanzee (Pan troglodytes), in a range of higher and lower order cortical areas, using a newly developed method that involves standard and immunohistochemical staining techniques to reveal and quantify the various profiles of the fine structures of the cerebral cortex. The results of these studies demonstrate clear layer differences in the wire fraction of the cerebral cortex, and for the most part, consistency in the neuropil wire fraction of the same layer across areas of the cerebral cortex within and between individuals of the same species; however, differences in the wire fraction of the neuropil were associated with changes in brain size. It is apparent that the neuropil is not static, as wiring “optimality” changes with layers and brain size and this has functional implications regarding neuronal processing and behavioural outcomes. The adaptive rationale adopted by evolutionary psychology studies to explain behaviours may be erroneous, as adaptation does not always explain sufficiently the emergence of complex behaviours related to brain size increases, especially in primates.

Sensory information is represented in cortex by cascades of excitation, the patterns of which are constrained and biased by anatomical connections between neurons. Additionally, in the living animal, functional connectivity is dynamically adjusted by internally generated background activity, which varies by arousal state and behavioral context. Therefore, to understand how excitation propagates through the cortex, it is necessary to characterize the laminar flow of signal propagation as well as spontaneous network activity, which will constrain that propagation. This thesis characterizes the nature and mechanisms of awake cortical network dynamics, as well as the sources of sensory inputs in different cortical layers of the rat somatosensory system. Mammalian brains generate internal activity independent of environmental stimuli. Internally generated states may bring about distinct cortical processing modes. To investigate how brain state impacts cortical circuitry, we recorded intracellularly from the same neurons, under anesthesia and subsequent wakefulness, in the rat barrel cortex. In every cell examined throughout layers 2-6, wakefulness produced a temporal pattern of synaptic inputs differing markedly from those under anesthesia. Recurring periods of synaptic quiescence, prominent under anesthesia, were abolished by wakefulness, which produced instead a persistently depolarized state. This switch in dynamics was unaffected by elimination of afferent synaptic input from thalamus, suggesting that arousal alters cortical dynamics by neuromodulators acting directly on cortex. Indeed, blockade of noradrenergic, but not cholinergic, pathways induced synaptic quiescence during wakefulness. This thesis shows that subcortical inputs from the locus coeruleus-noradrenergic system can switch local recurrent networks into different regimes via direct neuromodulation. Having characterized the nature of wakeful dynamics, I next sought to characterize how sensory information propagates through the cortex. The thalamocortical projection to layer 4 (L4) of primary sensory cortex is thought to be the main route by which information from sensory organs reaches the neocortex. Sensory information is believed to then propagate through the cortical column along the L4→L2/3→L5/6 pathway. This thesis shows that sensory-evoked responses of L5/6 neurons derive from direct thalamocortical synapses, rather than the intracortical pathway. A substantial proportion of L5/6 neurons exhibit sensory-evoked postsynaptic potentials and spikes with the same latencies as L4. Paired in vivo recordings from L5/6 neurons and thalamic neurons revealed significant convergence of direct thalamocortical synapses onto diverse types of infragranular neurons. Pharmacological inactivation of L4 had no effect on sensory-evoked synaptic input to L5/6 neurons, and responsive L5/6 neurons continued to discharge spikes. In contrast, inactivation of thalamus suppressed sensory-evoked responses. This thesis shows that L4 is not an obligatory distribution hub for cortical activity, contrary to long-standing belief, and that thalamus activates two separate, independent "strata" of cortex in parallel.

This thesis examines the dynamical behaviour of incorporating NMDA (an excitatory neurotransmitter) for the electrodynamic model of the cerebral cortex. The model used is the mean-field model developed by Steyn-Ross et al. (2005) which describes the behaviour of the cortex in terms of parameters averaged over spatially localised populations. The behaviour of the model is determined by the four control parameters: inhibitory effect li, subcortical drive s, and NMDA neurotransmitter e ect set by an excitatory factor le and the magnesium concentration C. Adopting this model could give a better understanding of the cortex functionality and the anaesthetic mechanism. The model predicts that there are either one or three stationary states available to the cortex. We identify two of these with highly activated state and a quiescent state and focus on the transition between the two. Theoretical stability predictions (eigenvalue analysis) verified by a numerical simulation show that the system is unstable between the two Hopf bifurcations. In addition, in the stable region the steady states remains stable under a small perturbation, while in the unstable region either a transition between states or a limit cycle (oscillation) occurs depending on the position of the steady state.

Thesis (Ph. D.)--Ohio State University, 2003. / Title from first page of PDF file. Document formatted into pages; contains xiii, 184 p.; also includes graphics (some col.). Includes abstract and vita. Advisor: Andrej Rotter, Dept. of Biochemistry. Includes bibliographical references (p. 158-184).

The posterior parietal cortex (PPC) is a cortical region receiving inputs from different sensory modalities which has been shown to subserve a visuospatial function. The potential contribution of PPC in audiospatial behaviours and recognition of amodal spatial correspondences were postulated and assessed in the present study. Adult male Long- Evans rats received PPC lesions by aspiration, and they were compared to sham operated control rats on three behavioural tasks. In the Morris water maze, the rats had to learn to use the distal visual cues to locate an escape platform hidden in the pool. In an open field task, the rats were assessed on their reactions to a spatial relocation of a visual or an auditory object. In a spatial cross-modal transfer (CMT) task (Tees & Buhrmann, 1989), rats were trained to respond to light signals using spatial rules, and were then subjected to transfer tests using comparable sound signals. Results from the Morris water maze, the open field, and the initial training phase of the spatial CMT task confirmed a visuospatial deficit in PPC lesioned rats. However, if given sufficient training, PPC lesioned rats could learn the location of a hidden platform in the Morris water maze, and they could also acquire spatial rules in the CMT task. Such results indicated that the visuospatial deficits in PPC lesioned rats were less severe than previously thought. On the other hand, a persistent navigational difficulty characterized by a looping pattern of movement was observed in the PPC lesioned rats in the Morris water maze. Results from the open field indicated that PPC was less involved in audiospatial behaviours. Moreover, results also indicated that PPC was not necessary for spatial CMT. Hence, data from the present study did not support the idea that PPC played an essential role in supramodal spatial abilities in the rats. Instead, data from the spatial CMT task seemed to imply a role of PPC in managing conflicting spatial information coming from different sensory modalities.

Thesis (Ph.D.)--Victoria University of Wellington, 2008. / Includes bibliographical references.

Thesis (Ph.D.)--University of Texas at Dallas, 2008. / Includes vita. Includes bibliographical references (leaves 105-106)