The brain is the part of the central nervous system that is contained in the cranial cavity of the skull. It includes the cerebral cortex, limbic system, basal ganglia, thalamus, hypothalamus, cerebellum, brainstem, and retinas. The outermost part of the brain is a thick piece of nervous system tissue called the cerebral cortex. The cerebral cortex, limbic system, and basal ganglia make up the two cerebral hemispheres. A thick fiber bundle called the corpus callosum (corpus = “body”; callosum = “tough”) connects the two hemispheres. Although there are some brain functions that are localized more to one hemisphere than the other, the functions of the two hemispheres are largely redundant. In fact, sometimes (very rarely) an entire hemisphere is removed to treat severe epilepsy. While patients do suffer some deficits following the surgery, they can have surprisingly few problems, especially when the surgery is performed on children who have very immature nervous systems.

In other surgeries to treat severe epilepsy, the corpus callosum is cut instead of removing an entire hemisphere. This causes a condition called split-brain, which gives insights into unique functions of the two hemispheres. For example, when an object is presented to patients’ left visual field, they may be unable to verbally name the object (and may claim to not have seen an object at all). This is because the visual input from the left visual field crosses and enters the right hemisphere and cannot then signal to the speech center, which generally is found in the left side of the brain. Remarkably, if a split-brain patient is asked to pick up a specific object out of a group of objects with the left hand, the patient will be able to do so but will still be unable to verbally identify it.

Each hemisphere contains regions called lobes that are involved in different functions. Each hemisphere of the mammalian cerebral cortex can be broken down into four functionally and spatially defined lobes: frontal, parietal, temporal, and occipital (Figure 16.22).

Figure 16.22 The human cerebral cortex includes the frontal, parietal, temporal, and occipital lobes.

The frontal lobe is located at the front of the brain, over the eyes. This lobe contains the olfactory bulb, which processes smells. The frontal lobe also contains the motor cortex, which is important for planning and implementing movement. Areas within the motor cortex map to different muscle groups. Neurons in the frontal lobe also control cognitive functions like maintaining attention, speech, and decision-making. Studies of humans who have damaged their frontal lobes show that parts of this area are involved in personality, socialization, and assessing risk. The parietal lobe is located at the top of the brain. Neurons in the parietal lobe are involved in speech and also reading. Two of the parietal lobe’s main functions are processing somatosensation—touch sensations like pressure, pain, heat, cold—and processing proprioception—the sense of how parts of the body are oriented in space. The parietal lobe contains a somatosensory map of the body similar to the motor cortex. The occipital lobe is located at the back of the brain. It is primarily involved in vision—seeing, recognizing, and identifying the visual world. The temporal lobe is located at the base of the brain and is primarily involved in processing and interpreting sounds. It also contains the hippocampus (named from the Greek for “seahorse,” which it resembles in shape) a structure that processes memory formation. The role of the hippocampus in memory was partially determined by studying one famous epileptic patient, HM, who had both sides of his hippocampus removed in an attempt to cure his epilepsy. His seizures went away, but he could no longer form new memories (although he could remember some facts from before his surgery and could learn new motor tasks).

Interconnected brain areas called the basal ganglia play important roles in movement control and posture. The basal ganglia also regulate motivation.

The thalamus acts as a gateway to and from the cortex. It receives sensory and motor inputs from the body and also receives feedback from the cortex. This feedback mechanism can modulate conscious awareness of sensory and motor inputs depending on the attention and arousal state of the animal. The thalamus helps regulate consciousness, arousal, and sleep states.

Below the thalamus is the hypothalamus. The hypothalamus controls the endocrine system by sending signals to the pituitary gland. Among other functions, the hypothalamus is the body’s thermostat—it makes sure the body temperature is kept at appropriate levels. Neurons within the hypothalamus also regulate circadian rhythms, sometimes called sleep cycles.

The limbic system is a connected set of structures that regulates emotion, as well as behaviors related to fear and motivation. It plays a role in memory formation and includes parts of the thalamus and hypothalamus as well as the hippocampus. One important structure within the limbic system is a temporal lobe structure called the amygdala. The two amygdala (one on each side) are important both for the sensation of fear and for recognizing fearful faces.

The cerebellum (cerebellum = “little brain”) sits at the base of the brain on top of the brainstem. The cerebellum controls balance and aids in coordinating movement and learning new motor tasks. The cerebellum of birds is large compared to other vertebrates because of the coordination required by flight.

The brainstem connects the rest of the brain with the spinal cord and regulates some of the most important and basic functions of the nervous system including breathing, swallowing, digestion, sleeping, walking, and sensory and motor information integration.

The limbic system is a connected set of structures that regulates emotion, as well as behaviors related to fear and motivation. It plays a role in memory formation and includes parts of the thalamus and hypothalamus as well as the hippocampus. One important structure within the limbic system is a temporal lobe structure called the amygdala (Greek for “almond”), illustrated in Figure 35.24. The two amygdala are important both for the sensation of fear and for recognizing fearful faces. The cingulate gyrus helps regulate emotions and pain.

Figure 35.24 The limbic system regulates emotion and other behaviors. It includes parts of the cerebral cortex located near the center of the brain, including the cingulate gyrus and the hippocampus as well as the thalamus, hypothalamus, and amygdala.

A set of forebrain structures common to all mammals that is defined functionally and anatomically. It is implicated in the higher integration of visceral, olfactory, and somatic information as well as homeostatic responses including fundamental survival behaviors (feeding, mating, emotion). For most authors, it includes the AMYGDALA; EPITHALAMUS; GYRUS CINGULI; hippocampal formation (see HIPPOCAMPUS); HYPOTHALAMUS; PARAHIPPOCAMPAL GYRUS; SEPTAL NUCLEI; anterior nuclear group of thalamus, and portions of the basal ganglia.

Other Areas of the Forebrain

Other areas of the forebrain, located beneath the cerebral cortex, include the thalamus and the limbic system. The thalamus is a sensory relay for the brain. All of our senses, with the exception of smell, are routed through the thalamus before being directed to other areas of the brain for processing (Figure).

The thalamus serves as the relay center of the brain where most senses are routed for processing.

Limbic System

The limbic system is involved in processing both emotion and memory. Interestingly, the sense of smell projects directly to the limbic system; therefore, not surprisingly, smell can evoke emotional responses in ways that other sensory modalities cannot. The limbic system is made up of a number of different structures, but three of the most important are the hippocampus, the amygdala, and the hypothalamus (Figure). The hippocampus is an essential structure for learning and memory. The amygdala is involved in our experience of emotion and in tying emotional meaning to our memories. The hypothalamus regulates a number of homeostatic processes, including the regulation of body temperature, appetite, and blood pressure. The hypothalamus also serves as an interface between the nervous system and the endocrine system and in the regulation of sexual motivation and behavior.

The Case of Henry Molaison (H.M.)

In 1953, Henry Gustav Molaison (H. M.) was a 27-year-old man who experienced severe seizures. In an attempt to control his seizures, H. M. underwent brain surgery to remove his hippocampus and amygdala. Following the surgery, H.M’s seizures became much less severe, but he also suffered some unexpected—and devastating—consequences of the surgery: he lost his ability to form many types of new memories. For example, he was unable to learn new facts, such as who was president of the United States. He was able to learn new skills, but afterward he had no recollection of learning them. For example, while he might learn to use a computer, he would have no conscious memory of ever having used one. He could not remember new faces, and he was unable to remember events, even immediately after they occurred. Researchers were fascinated by his experience, and he is considered one of the most studied cases in medical and psychological history (Hardt, Einarsson, & Nader, 2010; Squire, 2009). Indeed, his case has provided tremendous insight into the role that the hippocampus plays in the consolidation of new learning into explicit memory.

Forebrain Structures

The two hemispheres of the cerebral cortex are part of the forebrain (Figure), which is the largest part of the brain. The forebrain contains the cerebral cortex and a number of other structures that lie beneath the cortex (called subcortical structures): thalamus, hypothalamus, pituitary gland, and the limbic system (collection of structures). The cerebral cortex, which is the outer surface of the brain, is associated with higher level processes such as consciousness, thought, emotion, reasoning, language, and memory. Each cerebral hemisphere can be subdivided into four lobes, each associated with different functions.

Other Areas of the Forebrain

Other areas of the forebrain, located beneath the cerebral cortex, include the thalamus and the limbic system. The thalamus is a sensory relay for the brain. All of our senses, with the exception of smell, are routed through the thalamus before being directed to other areas of the brain for processing (Figure).

The four lobes of the brain are the frontal, parietal, temporal, and occipital lobes (Figure). The frontal lobe is located in the forward part of the brain, extending back to a fissure known as the central sulcus. The frontal lobe is involved in reasoning, motor control, emotion, and language. It contains the motor cortex, which is involved in planning and coordinating movement; the prefrontal cortex, which is responsible for higher-level cognitive functioning; and Broca’s area, which is essential for language production.

The lobes of the brain are shown.

People who suffer damage to Broca’s area have great difficulty producing language of any form (Figure). For example, Padma was an electrical engineer who was socially active and a caring, involved mother. About twenty years ago, she was in a car accident and suffered damage to her Broca’s area. She completely lost the ability to speak and form any kind of meaningful language. There is nothing wrong with her mouth or her vocal cords, but she is unable to produce words. She can follow directions but can’t respond verbally, and she can read but no longer write. She can do routine tasks like running to the market to buy milk, but she could not communicate verbally if a situation called for it.

Probably the most famous case of frontal lobe damage is that of a man by the name of Phineas Gage. On September 13, 1848, Gage (age 25) was working as a railroad foreman in Vermont. He and his crew were using an iron rod to tamp explosives down into a blasting hole to remove rock along the railway’s path. Unfortunately, the iron rod created a spark and caused the rod to explode out of the blasting hole, into Gage’s face, and through his skull (Figure). Although lying in a pool of his own blood with brain matter emerging from his head, Gage was conscious and able to get up, walk, and speak. But in the months following his accident, people noticed that his personality had changed. Many of his friends described him as no longer being himself. Before the accident, it was said that Gage was a well-mannered, soft-spoken man, but he began to behave in odd and inappropriate ways after the accident. Such changes in personality would be consistent with loss of impulse control—a frontal lobe function.

Beyond the damage to the frontal lobe itself, subsequent investigations into the rod's path also identified probable damage to pathways between the frontal lobe and other brain structures, including the limbic system. With connections between the planning functions of the frontal lobe and the emotional processes of the limbic system severed, Gage had difficulty controlling his emotional impulses.

However, there is some evidence suggesting that the dramatic changes in Gage’s personality were exaggerated and embellished. Gage's case occurred in the midst of a 19^th century debate over localization—regarding whether certain areas of the brain are associated with particular functions. On the basis of extremely limited information about Gage, the extent of his injury, and his life before and after the accident, scientists tended to find support for their own views, on whichever side of the debate they fell (Macmillan, 1999).

(a) Phineas Gage holds the iron rod that penetrated his skull in an 1848 railroad construction accident. (b) Gage’s prefrontal cortex was severely damaged in the left hemisphere. The rod entered Gage’s face on the left side, passed behind his eye, and exited through the top of his skull, before landing about 80 feet away. (credit a: modification of work by Jack and Beverly Wilgus)

The brain’s parietal lobe is located immediately behind the frontal lobe, and is involved in processing information from the body’s senses. It contains the somatosensory cortex, which is essential for processing sensory information from across the body, such as touch, temperature, and pain. The somatosensory cortex is organized topographically, which means that spatial relationships that exist in the body are maintained on the surface of the somatosensory cortex (Figure). For example, the portion of the cortex that processes sensory information from the hand is adjacent to the portion that processes information from the wrist.

Spatial relationships in the body are mirrored in the organization of the somatosensory cortex.

The temporal lobe is located on the side of the head (temporal means “near the temples”), and is associated with hearing, memory, emotion, and some aspects of language. The auditory cortex, the main area responsible for processing auditory information, is located within the temporal lobe. Wernicke’s area, important for speech comprehension, is also located here. Whereas individuals with damage to Broca’s area have difficulty producing language, those with damage to Wernicke’s area can produce sensible language, but they are unable to understand it (Figure).

Damage to either Broca’s area or Wernicke’s area can result in language deficits. The types of deficits are very different, however, depending on which area is affected.

The occipital lobe is located at the very back of the brain, and contains the primary visual cortex, which is responsible for interpreting incoming visual information. The occipital cortex is organized retinotopically, which means there is a close relationship between the position of an object in a person’s visual field and the position of that object’s representation on the cortex. You will learn much more about how visual information is processed in the occipital lobe when you study sensation and perception.

First, let's look at the role of the amygdala in memory formation. The main job of the amygdala is to regulate emotions, such as fear and aggression (Figure). The amygdala plays a part in how memories are stored because storage is influenced by stress hormones. For example, one researcher experimented with rats and the fear response (Josselyn, 2010). Using Pavlovian conditioning, a neutral tone was paired with a foot shock to the rats. This produced a fear memory in the rats. After being conditioned, each time they heard the tone, they would freeze (a defense response in rats), indicating a memory for the impending shock. Then the researchers induced cell death in neurons in the lateral amygdala, which is the specific area of the brain responsible for fear memories. They found the fear memory faded (became extinct). Because of its role in processing emotional information, the amygdala is also involved in memory consolidation: the process of transferring new learning into long-term memory. The amygdala seems to facilitate encoding memories at a deeper level when the event is emotionally arousing.

The amygdala is involved in fear and fear memories. The hippocampus is associated with declarative and episodic memory as well as recognition memory. The cerebellum plays a role in processing procedural memories, such as how to play the piano. The prefrontal cortex appears to be involved in remembering semantic tasks.

The amygdala is a group of nuclei in the medial region of the temporal lobe that is part of the limbic lobe (Figure 15.12). The limbic lobe includes structures that are involved in emotional responses, as well as structures that contribute to memory function. The limbic lobe has strong connections with the hypothalamus and influences the state of its activity on the basis of emotional state. For example, when you are anxious or scared, the amygdala will send signals to the hypothalamus along the medial forebrain bundle that will stimulate the sympathetic fight-or-flight response. The hypothalamus will also stimulate the release of stress hormones through its control of the endocrine system in response to amygdala input.

Figure 15.12 The Limbic Lobe Structures arranged around the edge of the cerebrum constitute the limbic lobe, which includes the amygdala, hippocampus, and cingulate gyrus, and connects to the hypothalamus.