This paper is a rough precis of a recent book (GRAY, 1982). It is concerned with the question: what are the brain structures which mediate the psychology as well as the neurology of anxiety? It is clear that, once you start asking questions about the brain, you have to work with animals for reasons that are familiar to everybody. But it is very difficult to start studying emotions in animals, because it is difficult to know what emotions animals experience and whether these are the same as the human emotions. So the first thing one has to look for is a kind of crutch or bridge to get you across from the human domain on one side to the animal domain on the other. Now in the case of anxiety we are quite fortunate in that there are certain drugs which are clinically very effective in controlling anxiety in human subjects. So one can take those drugs and apply them to animals to study their behavioral effects.

https://link.springer.com/content/pdf/10.1007%2F978-3-642-69631-2_5.pdf

The neuropsychology of anxiety

The purpose of the present investigation was to examine the topographical organization of efferent projections from the cytoarchitectonic divisions of the mPFC (the medial precentral, dorsal anterior cingulate and prelimbic cortices). We also sought to determine whether the efferents from different regions within the prelimbic division were organized topographically. Anterograde transport of Phaseolus vulgaris leucoagglutinin was used to examine the efferent projections from restricted injection sites within the mPFC. Major targets of the prelimbic area were found to include prefrontal, cingulate, and perirhinal cortical structures, the dorsomedial and ventral striatum, basal forebrain nuclei, basolateral amygdala, lateral hypothalamus, mediodorsal, midline and intralaminar thalamic nuclei, periaqueductal gray region, ventral midbrain tegmentum, laterodorsal tegmental nucleus, and raphe nuclei. Previously unreported projections of the prelimbic region were also observed, including efferents to the anterior olfactory nucleus, the piriform cortex, and the pedunculopontine tegmental-cuneiform region. A topographical organization governed the efferent projections from the prelimbic area, such that the position of terminal fields within target structures was determined by the rostrocaudal, dorsoventral, and mediolateral placement of the injection sites. Efferent projections from the medial precentral and dorsal anterior cingulate divisions (dorsomedial PFC) were organized in a similar topographical fashion and produced a pattern of anterograde labeling different from that seen with prelimbic injection sites. Target structures innervated primarily by the dorsomedial PFC included certain neocortical fields (the motor, somatosensory, and visual cortices), the dorsolateral striatum, superior colliculus, deep mesencephalic nucleus, and the pontine and medullary reticular formation. Previously unreported projections to the paraoculomotor central gray area and the mesencephalic trigeminal nucleus were observed following dorsomedial PFC injections. These results indicate that the efferent projections of the mPFC are topographically organized within and across the cytoarchitectonic divisions of the medial wall cortex. The significance of topographically organized and restricted projections of the rat mPFC is discussed in light of behavioral and physiological studies indicating functional heterogeneity of this region.

Topographical organization of the efferent projections of the medial prefrontal cortex in the rat: An anterograde tract-tracing study with Phaseolus vulgaris leucoagglutinin

Functions of the frontal cortex of the rat: A comparative review

The study of memory is a great challenge, perhaps the greatest in biological sciences. Memory involves changes in a tiny fraction of an extremely large pool of elements, a conclusion that makes the task of finding those changes using current technologies formidable. What can be done about this roadblock to neurological investigations of learning? One response that has become particularly productive in recent years is to study learning or learning-like phenomena in relatively simple "model" systems. The idea is to extract basic principles from these models in which molecular and anatomical details can be studied and then to use these in analyzing learning in higher regions of the brain. In this article we discuss current progress and emerging concepts derived from the simple system approach using animal models.

The neurobiology of learning and memory

The connections between the olfactory bulb, primary olfactory cortex, and olfactory related areas of the orbital cortex were defined in macaque monkeys with a combination of anterograde and retrograde axonal tracers and electrophysiological recording. Anterograde tracers placed into the olfactory bulb labeled axons in eight primary olfactory cortical areas: the anterior olfactory nucleus, piriform cortex, ventral tenia tecta, olfactory tubercle, anterior cortical nucleus of the amygdala, periamygdaloid cortex, and olfactory division of the entorhinal cortex. The bulbar axons terminate in the outer part of layer I throughout these areas and are most dense in areas that are close to the lateral olfactory tract. Labeled axons also were found in the superficial part of nucleus of the horizontal diagonal band. Retrograde tracers injected into the olfactory bulb labeled cells in the nucleus of the diagonal band and in all of the primary olfactory cortical areas except the olfactory tubercle. Electrical stimulation of the olfactory bulb evoked short-latency unit responses and a characteristic field wave in the primary olfactory cortex. Multiunit activity in layer II tended to be of shorter latency than that in layer III and the endopiriform nucleus. Associational connections within the primary olfactory cortex were demonstrated with anterograde tracer injections into the piriform cortex and the entorhinal cortex. Injections into the piriform cortex near the lateral olfactory tract labeled axons in the deep part of layer I of many primary olfactory areas, but especially in areas near the tract. An injection into the rostral entorhinal cortex, distant to the lateral olfactory tract, labeled a complementary distribution of axons in deep layer I of olfactory areas medial and caudoventral to the tract. This organization resembles that reported in the primary olfactory cortex of the rat [Luskin and Price (1983) J. Comp. Neurol. 216:264-291]. The anterograde tracer injections into the piriform cortex and retrograde tracer injections into the orbital and medial prefrontal cortex and rostral insula label connections from the primary olfactory cortex to nine areas in the caudal orbital cortex, including the agranular insula areas Iam, Iai, Ial, Iapm, and Iapl and areas 14c, 25, 13a, and 13m. The piriform cortex projects most heavily to layer I of these areas. Only Iam, Iapm, and 13a receive a substantial projection to the deeper layers. Areas Iam, Iapm, and 13a were also the only areas that responded with multiunit action potentials to olfactory bulb stimulation in anesthetized animals.(ABSTRACT TRUNCATED AT 400 WORDS)

Central Olfactory Connections in the Macaque Monkey

The "core" subnucleus of the mediodorsal thalamic nucleus (MD) receives direct input from olfactory cortex in the rat. This part of MD projects to the frontal neocortex of the rhinal sulcus (RS), while other parts of the MD project to the anterior medial wall of the neocortex (MW). In this study, rats were tested on odor threshold, detection of specific odors, and three odor discriminations both before and after lesions of MD, MW, RS, or sham lesions. Olfactory threshold and detection ability remained unaffected by any of these lesions; however, subjects with MD and RS lesions were markedly impaired on odor discriminations. Furthermore, subjects were more profoundly impaired when the stimuli to be discriminated were novel or difficult to associate. Thus, MD and its "olfactory" projection target, RS, seem to mediate cognitive, rather than sensory aspects of odor-guided behavior in the rat.

Thalamocortical mechanisms in odor-guided behavior. I. Effects of lesions of the mediodorsal thalamic nucleus and frontal cortex on olfactory discrimination in the rat.

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Thalamocortical mechanisms in odor-guided behavior. I. Effects of lesions of the mediodorsal thalamic nucleus and frontal cortex on olfactory discrimination in the rat.