Review the functions of the limbic system. Explain its role in the control of major affective activities.

Dionne Angela Donnelly Module Code: PSYC121 Tutorial Essay Year One

The limbic system (LS) is a collection of nuclei and cortical brain structures found underneath the neocortex, within the deep medial sections in the forebrain (Isaacson, 1982). It is thought to include five major structures: the limbic cortex, amygdala, hippocampus, fornix and mammillary bodies (Carlson, 2007), which are all linked by their location and functions. These structures are involved in learning and memory, and a range of affective activities (i.e. behavioural, cognitive or biological activities which relate to emotion), such as feelings and expressions of emotions, emotional memories, and recognition of signs of emotion in others (e.g. Panksepp, 2004). However, the list of structures noted above is by no means conclusive. There is much debate as to which structures should be included or omitted. There has also been speculation as to whether the term should either face a change in definition or become obsolete (Ktter & Stephan, 1997). Also, the focus here will only be upon the five structures noted above, as these appear to be those which are least controversial within the neuroscientific community.

It was Papez (1937, cited in Carlson, 2007) who first stated that a single set of brain structures were responsible for human affect and motivation. He termed this the 'Papez circuit'. This circuit can be found at the core of the forebrain, in front of the brainstem. It consists of nerve fibres which connect the thalamus, hypothalamus, anterior thalamus, cingulate cortex and hippocampus into a 'loop' (Cornelius, 1996). Papez (1937, cited in Strongman, 2006) believed that this circuit played the most important role in the regulation and expression of our emotions, and that this expression of emotions (i.e. behavioural affective activities such as smiling, crying etc.) was dependent on the hypothalamus, whereas feelings or the experience of emotion rely upon the cortex. One criticism of this theory is that he did not include other brain structures which have later been found to play a role in affective activities, such as the primary cortex. A second limitation is that it is quite vague as to how each structure behaves and what emotions, if any, they each relate to. Despite this, the theory has led the way for further research into the emotional aspects of the LS.

This further research was conducted by MacLean (e.g. 1949, 1970). He adapted the Papez circuit to include the amygdala, fornix and mammillary bodies (MacLean, 1949, cited in Carlson, 2007). This 'control centre of emotion' is called the LS, although MacLean clearly placed less emphasis on the hypothalamus than Papez, as it does not share in the experiencing of emotion (Masserman, 1943, cited in MacLean, 1949, p. 338). MacLean (1970, cited in Cornelius, 1996) posited that the 'effector system' is most likely to be the hypothalamus, but it is the LS which integrates emotional information received via its numerous subcortical connections. Cornelius (1996) argued that the most important structures within the LS are the hippocampus and the amygdala, which are intricately connected with one another and both receive signals from the senses and key organs within the body that we associate with emotion, such as the heart and the intestines (or 'gut'). Cornelius (1996) also states that it is the role of the LS to integrate these signals with information received from the autonomic nervous system (ANS, i.e. part of the nervous system which regulates involuntary functions such as heartbeat and respiration).

LeDoux (1995) postulated that the idea that the LS is the sole structure involved in affective activities has fallen into disfavour, along with the concept that the LS is even a discrete anatomical structure (LeDoux, 1995, cited in Cornelius, 1996). Therefore there must be other brain structures which play a role in our affective activities. Researchers such as Panksepp (e.g. 1994) have argued that the LS is only one part of a wider circuit which relates to affective activities. This circuit includes interaction between the midbrain, the LS and the basal ganglia. These are said to be linked by four emotion-mediating circuits relating to expectancy, fear, rage, and panic. Bluhm (2008) argued that the 'uniqueness' of human affect is reliant upon the interaction between the prefrontal cortex, cognition and LS. Therefore, although the LS clearly plays an important role in affective activities, it is also clear that other brain areas/structures have a profound effect upon how we behave emotionally. This puts the debate over whether the LS should still be used as a concept into the spotlight, as the LS is defined by the fact that the structures are linked by the function of
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emotion, but it seems that most of the brain plays some role in this. Watt (1998) states the borders of the limbic system are vague and have been extended decade by decade like the erosion of a vast neural shoreline. This raises a number of questions: should more areas be included in the concept of the LS? Or should the term be scrapped? Ktter & Stephan (1997) argue that the LS is used to describe too many functions, it has become an all-encompassing concept which is vague and in some cases contradictory, and it is therefore difficult to interpret the true functions of the structures it contains.

As the descriptions above only offer only a view of the LS as a whole, it is necessary, in order to attempt to fully understand the numerous functions, to not only look at the complete system but also each structure that comprises it in isolation, as this will provide a more in-depth view of the functions of each part and will make clear each structure's role in human affective activities.

The limbic cortex is the border which surround the LS, it contains the cingulate gyrus which is just above the corpus callosum. The cingulate gyrus has been found to be involved in error related negativity, self-esteem, depression, decision making and emotional processing (Fragopanagos & Taylor, 2007). Papez noted that tumours within the limbic cortex are associated with changes of personality, loss of affect, and various degrees of somnolence and stupor (MacLean, 1949, p. 341). More recent research has discovered damage to the anterior cingulate cortex (ACC) can not only lessen emotional experience, but can also completely eradicate it (Damasio & Van Hoesen, 1983, cited in Prinz, 2006), this is known as akinetic mutism. The cingulate gyrus has also been associated with sexuality. One study followed a hypersexual 55 year-old female who developed convulsions exacerbated by perfumes. It was discovered that she had a hemangioma (a benign tumorous mass of blood vessels) just above the cingulate gyrus (Erikson, 1945, cited in MacLean, 1949). The ACC also receives signals from other cortical areas e.g. the insula, this means the ACC may be the focus of intermediate-level emotional processing (Prinz, 2006, p. 155). This evidence
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clearly shows that whilst the limbic cortex/cingulate gyrus has an important role in affective activities, it is involved in other aspects of behaviour and is reliant on input from other areas of the brain.

One of the most important structures to consider in relation to affective activities is the amygdala. Research has determined that the amygdala serves several crucial functions, not only affecting emotional behaviour and motivation, but also serving memory function (Reber & Reber, 2001, p. 30). The amygdala is an almond shaped group of nuclei, found bilaterally inside the anterior temporal lobe (Isaacson, 1974, cited in Cornelius, 1996). It is most commonly noted for its role in fear, which may relate to the evolution of the LS in mammals as a survival function (the LS is also referred to as the paleopallium/old mammalian brain). For example, if the amygdala is removed in animals, they show no response to situations which usually invoke stress and anxiety, meaning they cannot successfully survive in the wild (Carlson, 2004), especially since this may affect their 'fight or flight' response. Another function of the amygdala appears to be in the response to taste, as it is part of the anteromedial temporal lobe (AMTL), when this is removed, people have an increased sensitivity to bitter tastes (Small et al, 1997, cited in Carlson, 2004). When the amygdala is stimulated via electricity, chemicals or drugs, animals display physiological and behavioural aspects of fear and agitation (Carlson, 2004). When the amygdala is damaged in humans they lose the ability to recognise other people's fear expressions, cannot draw fearful expressions and have difficulty recognising sounds related to fear (Carlson, 2004). Bauman and Kemper (1994, cited in Carlson, 2004) have found that in some cases of autism (a syndrome in which people cannot recognise emotion in others and have trouble coping with social situations), small, closely situated neurons appear in the amygdala and the hippocampus, suggesting they both play a role in our understanding of the emotions of others.

The amygdala therefore obviously has an important role in recognising emotion and in regulating the fear response. However, as the amygdala appears to be so specialised, it does imply that other areas of the brain must be involved in other affective activities (i.e. the expression of happiness, sadness etc.). It has also been noted that different areas of the amygdala may be responsible for different activities. For example, the medial nucleus receives information from the senses regarding olfaction (Carlson, 2007). This is important because the LS is the area which was referred to as the rhinencephalon or 'smell brain'. It has been noted that the majority of those structures involved in emotional experience have also been linked to olfactory function, but this was refuted by Papez (1937, cited in MacLean, 1949). He stated that there was clinical or other evidence to support this view. However, this is debatable because another structure of the LS, which is closely linked to the amygdala, has also been linked olfaction.

The hippocampus, like all structures within the LS, has a number of functions. It is located in the temporal lobe beside the amygdala, hence their close relationship. Research by Deshmukh and Bhalla (2003) has found that different neurons within the hippocampus fire depending upon different odours presented. Levy, Hopkins and Squire (2004) also found that odour recognition span was greatly decreased by lesions to the hippocampal region. Therefore, it seems that the LS does seem to perform olfactory functions as well as affective functions. The hippocampus has also been linked to memory. Vargha- Khadem, Gadian, Watkins, Connelly, Van Paesschen, and Mishkin (1997, cited in Carlson, 2007) state that when the hippocampus is destroyed, episodic memory is affected, and in order to prevent the storage of all declarative memories, the hippocampus and limbic cortex would have to be destroyed/removed. The hippocampus is also associated with spatial awareness and memory. Luzzi, Pucci, Bella and Piccirilli (2000, cited in Carlson, 2007) studied a patient with a lesion to the right parahippocampal gyrus. This patient found it impossible to navigate around new environments and had to use cues such as the number of doors to find his own room. In relation to affective activities the hippocampus appears to be specifically important in our
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response to novel stimuli (e.g. surprise). Knight (1996, cited in Carlson, 2004) found that patients with damage to the hippocampus, physiological responses to novel stimuli were significantly less than those of normal controls. A further finding was that both groups reacted in the same way to stimuli that were expected, which suggests that the hippocampus has a role in emotion for survival, as it guides the individual towards new stimuli that whilst maybe alarming, could prevent serious harm to the individual. Consequently, it appears that the hippocampal role in affective activities is to supplement the activities of the amygdala and to integrate emotional experience into memory.

However, the hippocampus is not only linked to the amygdala, it is also connected via the fornix to the mammillary bodies of the posterior hypothalamus. The fornix has been found to be linked to fear conditioning. Phillips and LeDoux (1995) found that lesions of the fornix disrupted fear conditioning in rats, which suggests that the lesion disrupted the connection between the hypothalamus and the hippocampus/amygdala, as these are the areas associated with fear. Kuzniecky, Bilir, Gilliam, Faught, Martin, and Hugg (1999) studied epilepsy patients using MRI and found that areas of the LS, in particular the fornix, were more likely to be atrophied than control subjects. This suggests that the fornix also has a functional role in epilepsy. The mammillary bodies appear to play a role in emotion and memory, specifically involved in the alcohol-related WernickeKorsakoff syndrome, as it was found that experimental lesions of the mammillary bodies had similar detrimental effects on memory as damage due to thiamine deficiency as a result of chronic alcohol consumption (CAC) (Bracocha, 2005). This damage also led to the inability to remain emotionally aroused for long periods of time, thus suggesting that the mammillary bodies play an extremely important role in the regulation of our affective activities, as they affect our ability to not only feel emotion but to display emotional behaviours.

Despite debate about whether the LS should still be considered as a valid, functional concept, it appears that the structures of the LS do play a major role in relation to affective activities. Although
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they interact with other areas of the brain such as the prefrontal cortex, it is the structures of the LS which appear to perform the crucial functions in regulating our affective activities. Also, the separate structures if the LS have a range of functions, such as taste, memory and olfaction, which all seem to interact with one another to mediate each individuals affective response to their environment.

References Bracocha, D. (2005). Interaction Between Emotion and Memory: Importance of Mammillary Bodies Damage in a Mouse Model of the Alcoholic Korsakoff Syndrome. Neural Plasticity, 12 (4), 275-287. Bhalla, U. S., & Deshmuhk, S. S. (2003). Representation of Odor Habituation and Timing in the Hippocampus. The Journal of Neuroscience, 23(5), 1903. Bilir, E., Gilliam, f., Faught, E., Hugg, J., Kuzniecky, R., & Martin, R. (1999). Quantitative MRI in Temporal Lobe Epilepsy. Neurology, 53, 496. Bluhm, R. (2006). Beyond the Basics. The Modularity of Emotions. Canadian Journal of Philosophy, 32 (supplementary) 73-94. Canada: University of Calgary Press. Buskist, W., Carlson, N. R., Martin, G. N. (2004). Psychology (2nd Ed.). UK: Pearson Education. Carlson, N. R. (2007). Physiology of Behaviour (9th Edition). USA: Pearson Education. Cornelius, R. R. (1996). The Science of Emotion. Research and Tradition in the Psychology of Emotion. New Jersey: Prentice-Hall. Hopkins, R. O., Levy, D. A., & Squire, L. R., (2004). Impaired Odor Recognition Memory in Patients With Hippocampal Lesions. Learning & Memory 11, 794-796. Isaacson, R. L. (1982). The Limbic System. New York: Plenum Ktter, R., & Stephan, K., E. (1997). Useless or Helpful? The Concept Limbic System. Reviews in the Neurosciences, 8, 139-145.

Ledoux, J. E. & Philips, R. G. (1995). Lesions of the Fornix but not the Entorhinal or Perirhinal Cortex Interfere with Contextual Fear Conditioning. The Journal of Neuroscience, 15 (7), 53085315. MacLean, P. D. (1949) Psychosomatic Disease and the "Visceral Brain" Recent Developments Bearing on the Papez Theory of Emotion. Psychosomatic Medicine 11, 338-353 Panksepp, J. (1994). Affective Neuroscience The Foundations of Human and Animal Emotions. USA: Oxford University Press. Prinz, J. J. (2006). Is Emotion a Form of Perception? The Modularity of Emotions. Canadian Journal of Philosophy, 32 (supplementary) 137-160. Canada: University of Calgary Press. Reber, A. S., & Reber, E. (2001). Dictionary of Psychology (3rd Ed.).London: Penguin Books Strongman, K. T. (2006) The Psychology of Emotion From Everyday Life To Theory (5th Edition). West Sussex: John Wiley & Sons. Watt, D. F. (1998). Affect and the Limbic System: Some Hard Questions. Journal of Neuropsychiatry & Clinical Neuroscience 10:113-116.