The Papez circuit , or medial limbic circuit, is a neural circuit for the control of emotional expression. In 1937, James Papez proposed that the circuit connecting the hypothalamus to the limbic lobe was the basis for emotional experiences. Paul D. MacLean reconceptualized Papez's proposal and coined the term limbic system. MacLean redefined the circuit as the "visceral brain" which consisted of the limbic lobe and its major connections in the forebrain – hypothalamus, amygdala, and septum. Over time, the concept of a forebrain circuit for the control of emotional expression has been modified to include the prefrontal cortex.
Structure
The Papez circuit involves various structures of the brain. It begins and ends with the hippocampus (or the hippocampal formation). Fiber dissection indicates that the average size of the circuit is 350 millimeters. The Papez circuit goes through the following neural pathways:
Hippocampal formation (subiculum) → fornix → mammillary bodies → mammillothalamic tract → anterior thalamic nucleus → cingulum → entorhinal cortex → hippocampal formation. A photograph of the inferior medial view of the brain when dissected clearly shows the layout of the Papez circuit. Due to the location of the structures in the circuit, the resulting shape is a limbus. This is what drove MacLean to call the circuit the limbic system when he later modified the circuit. Various studies indicate that the Papez circuit is greatly influenced by the cerebellum and that perhaps the hippocampus is not the starting point of the circuit. Anatomically, this would make sense since the cerebellum is connected to the circuit with many fine fibers and fiber bundles. Chemical lesions on the cerebellum seem to have an inhibitory effect on the circuit. Animal behavioral studies show that electrical stimulation of the anterior cerebellum can cause arousal, predatory attack, and feeding responses, all of which are thought to be expressions of emotion. Overall, these studies provide evidence that the cerebellum may also be included in an emotional system of the brain. In recent years, multiple additional limbic fiber connectivity has been revealed using Diffusion MRI-weighted imaging MRI techniques. The equivalent fiber connectivity of all these pathways has been documented by dissection studies in primates. The fiber tracts include the amygdalofugal tract, amygdalothalamic tract, stria terminalis, dorsal thalamo-hypothalamic tract, cerebellohypothalamic tracts, and the parieto-occipito-hypothalamic tract.
Function
Emotion
Based on Papez's experiment with aggression in rats and other studies, it was initially believed that the circuit was involved with emotion. The circuit connects the hypothalamus and the cortex and acts as the emotional system of the brain. "The cingulate cortex projects to the hippocampus, and the hippocampus projects to the hypothalamus by way of the bundle of axons called the fornix. Hypothalamic effects reach the cortex via a relay in the anterior thalamic nuclei." However, there has not been additional proof of the circuit's role in emotion over the years. Now, the Amygdala is thought to play a key role in emotion, a structure that was not a part of the Papez circuit until 1952 when MacLean included it in the modified version of the circuit, the limbic system.
Memory In the past few years, many experiments have been done that involve structures of the Papez circuit. Moreover, many cases where structures of the Papez circuit have been damaged show changes that indicate its real function. Theta waves are used to measure activity in mainly the hippocampus but can be used to measure other brain regions as well. Strong synchronization of theta waves in the hippocampus and anterior ventral thalamus has been observed. Although theta waves between the hippocampus and the anterior dorsal and anterior medial regions in the thalamus showed no synchronization, scientists have seen a marked increase in firing rates of the neurons in the brain regions. Overall, based on theta wave experimentation, some components of the Papez circuit have shown to indeed have a connection and work together. Also, theta waves are thought to be linked to learning and memory. Consequently, many scientists believe that the Papez circuit is involved with memory because of theta wave experiments. Some scientists have narrowed it down to specifically spatial and episodic memory. Damage to the mammillothalamic tract, ventral anterior nucleus, and ventral lateral nucleus can result in memory and language impairment. Amnesia can be a result of disconnection of the mammillary bodies from the Papez circuit. The fornix is a bundle of nerve tracts made of white matter. It is crucial in normal cognitive functions. Harm to the fornix can result in amnesia.
Clinical significance
Alzheimer's disease Alzheimer's disease is a common neurodegenerative disease and a form of dementia. It leaves patients impaired or unable to learn new information. They usually have difficulty coming up with the names of people, objects, etc. Degeneration or other neural problems because of this disease commonly occurs in parts of the Papez circuit. It is already known that problems with episodic memory are linked to tumors or damage in the Papez circuit. Specific structures of the Papez circuit linked to episodic memory are the hippocampus and other medial temporal lobe structures. As a result of these adverse effects on episodic memory, damage to the Papez circuit can not only indicate or predict amnesia but also Alzheimer's in a patient.
Parkinson's disease Parkinson's disease is caused by the degeneration of the substantia nigra in the midbrain. There may also be damage to the structures of the Papez circuit in this disease. Usually, Parkinson's is correlated with depletion of dopamine in the basal ganglia.
Semantic dementia Semantic dementia is a rare degenerative disorder that exhibits defects in all semantic memory functions, including naming, single word comprehension and impoverished general knowledge, with relative preservation of other components of speech, perceptual and nonverbal problem-solving skills, and episodic memory. This may occur due to damage in the mammillary bodies, ventral anterior nucleus, and ventral lateral nucleus which has resulted in disruption in language in previous research.
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