Stimulation or lesions in other regions of the limbic system, especially in the amygdala, the septal area, and

areas in the mesencephalon, often cause effects similar to those elicited from the hypothalamus. We discuss
some of these in more detail later.

“Reward” and “Punishment” Function of the Limbic System

Electrical stimulation of certain limbic areas pleases or satisfies the animal, whereas electrical stimulation of other
regions causes terror, pain, fear, defense, escape reactions, and all the other elements of punishment (the reward and
punishment centers). The degrees of stimulation of these two oppositely responding systems greatly affect the
behavior of the animal.

Importance of Reward or Punishment on Behavior

Almost everything that we do is related in some way to reward and punishment. If we are doing something that is
rewarding, we continue to do it; if it is punishing, we cease to do it. Therefore, the reward and punishment centers
undoubtedly constitute one of the most important of all the controllers of our bodily activities, our drives, our
aversions, our motivations.

Effect of Tranquilizers on the Reward or Punishment Centers.

Administration of a tranquilizer, such as chlorpromazine, usually inhibits both the reward and the punishment
centers, thereby decreasing the affective reactivity of the animal. Therefore, it is presumed that tranquilizers function
in psychotic states by suppressing many of the important behavioral areas of the hypothalamus and its associated
regions of the limbic brain.

Specific Functions of Other Parts of the Limbic System Functions of the Hippocampus

The hippocampus is the elongated portion of the cerebral cortex that folds inward to form the ventral surface
of much of the inside of the lateral ventricle. One end of the hippocampus abuts the amygdaloid nuclei, and along its
lateral border it fuses with the parahippocampal gyrus, which is the cerebral cortex on the ventromedial outside
surface of the temporal lobe.

The hippocampus (and its adjacent temporal and parietal lobe structures, all together called the hippocampal
formation) has numerous but mainly indirect connections with many portions of the cerebral cortex, as well as with
the basal structures of the limbic system—the amygdala, hypothalamus, septum, and mamillary bodies. Almost any
type of sensory experience causes activation of at least some part of the hippocampus, and the hippocampus in turn
distributes many outgoing signals to the anterior thalamus, hypothalamus, and other parts of the limbic system,
especially through the fornix, a major communicating pathway. Thus, the hippocampus is an additional channel
through which incoming sensory signals can initiate behavioral reactions for different purposes. As in other limbic
structures, stimulation of different areas in the hippocampus can cause almost any of the different behavioral
patterns such as pleasure, rage, passivity, or excess sex drive.
Another feature of the hippocampus is that it can become hyperexcitable. For instance, weak electrical stimuli can
cause focal epileptic seizures in small areas of the hippocampi. These often persist for many seconds after the
stimulation is over, suggesting that the hippocampi can perhaps give off prolonged output signals even under normal
functioning conditions. During hippocampal seizures, the person experiences various psychomotor effects, including
olfactory, visual, auditory, tactile, and other types of hallucinations that cannot be suppressed as long as the seizure
persists even though the person has not lost consciousness and knows these hallucinations to be unreal. Probably one
of the reasons for this hyperexcitability of the hippocampi is that they have a different type of cortex from that
elsewhere in the cerebrum, with only three nerve cell layers in some of its areas instead of the six layers found
elsewhere.

Role of the Hippocampus in Learning

Effect of Bilateral Removal of the Hippocampi— Inability to Learn.

Portions of the hippocampi have been surgically removed bilaterally in a few human beings for treatment of
epilepsy. These people can recall most previously learned memories satisfactorily. However, they often can learn
essentially no new information that is based on verbal symbolism. In fact, they often cannot even learn the names of
people with whom they come in contact every day. Yet they can remember for a moment or so what transpires
during the course of their activities. Thus, they are capable of short-term memory for seconds up to a minute or two,
although their ability to establish memories lasting longer than a few minutes is either completely or almost
completely abolished. This is the phenomenon called anterograde amnesia

Functions of the Amygdala

The amygdala is a complex of multiple small nuclei located immediately beneath the cerebral cortex of the medial anterior pole
of each temporal lobe. It has abundant bidirectional connections with the hypothalamus, as well as with other areas of the limbic
system.

In lower animals, the amygdala is concerned to a great extent with olfactory stimuli and their interrelations with the limbic brain.
one of the major divisions of the olfactory tract terminates in a portion of the amygdala called the corticomedial nuclei, which
lies immediately beneath the cerebral cortex in the olfactory pyriform area of the temporal lobe. In the human being, another
portion of the amygdala, the basolateral nuclei, has become much more highly developed than the olfactory portion and plays
important roles in many behavioral activities not generally associated with olfactory stimuli.

The amygdala receives neuronal signals from all portions of the limbic cortex, as well as from the neocortex of the temporal,
parietal, and occipital lobes—especially from the auditory and visual association areas. Because of these multiple connections,
the amygdala has been called the “window” through which the limbic system sees the place of