TWO

Pride and Prejudice and Brains

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THROUGH DARCY’S EYES

Let’s start with a look, a catching of the eye. Vision is, after all, the sense that most of us rely on for information about the world. So we’ll begin with the look exchanged between Darcy and Elizabeth at the Meryton ball, where they first meet.

The ball is a thrilling event for Meryton’s excited young ladies and their hopeful mothers since the community has recently acquired two eligible bachelors, Darcy and Bingley. Their presence transforms the ball from a country dance to a matter of life and death—or at least of financial solvency—since in Austen’s day, marriage was the only respectable career option for daughters of upper- and middle-class families. Those who lacked family money and failed to marry faced humiliating and potentially harsh options: They would either have to depend on the charity of friends and family, or take a position as a paid companion or governess. (The latter is a prospect dreaded by Jane Fairfax in Emma.) To further complicate things for the Bennet family, they have no male heirs, and so the family estate is destined to go to the closest male relative. The Bennets would have no other residence or source of income upon the death of Mr. Bennet, and so the family needs to marry off all five of its daughters.*

As we discussed in the previous chapter, Bingley enjoys himself immensely at the ball, in part because he’s falling in love with Elizabeth’s sister Jane. But Darcy stiffly endures the event from the sidelines. When Bingley encourages Darcy to dance, he refuses, declaring that there’s no one in the room worth dancing with. Bingley gestures toward Jane’s sister Elizabeth, who’s sitting close enough to overhear their conversation, but Darcy isn’t convinced. He looks in her direction and scornfully replies,

Which [who] do you mean?” and turning round he looked for a moment at Elizabeth, till catching her eye he withdrew his own and coldly said, “She is tolerable; but not handsome enough to tempt me; and I am in no humour at present to give consequence to young ladies who are slighted by other men. You had better return to your partner and enjoy her smiles, for you are wasting your time with me. [my italics]

Darcy’s contemptuous glance begins what well might be the most famous romance in all of English literature. But let’s consider it from a more technical point of view, in terms of what actually happens in Darcy’s brain when he looks at Elizabeth.

This point of view might not come naturally, or even be all that enticing, to literature buffs. As a society, our education and our interests all too often divide between the arts and the sciences. But I’m convinced that Austen would have appreciated knowing about the brain. Although she wasn’t a scientist, she was nevertheless a most astute observer of human nature along what we recognize as scientific principles. She never wrote about things she couldn’t witness, such as men’s conversations when alone. Equipped with astute powers of observation, she came to understand people so accurately that we can use her novels to illustrate the science of social intelligence.

Austen would have realized that psychological understanding is deepened and expanded by knowing about the brain as well as the mind. Today, most psychologists no longer think in terms of a mind vs. a brain, but of a mind-brain. And the different disciplines that focus on one or the other of these entities have come to be so intertwined that their boundaries are no longer distinct. Psychologists understand the basics of neuroscience, and neuroscientists explore theories that used to be considered purely psychological. Researchers often identify with more than one of these fields.

This dual focus makes sense because the brain is an important part of the mind. In fact, most mind-brain scientists maintain that the mind and brain are actually the same, and that the difference between the two pertains to an observer’s perspective rather than to the mind-brain in and of itself. They believe that they’re two sides of a coin, and that the mind arises from the activity of the brain. However, I follow psychiatrist Daniel Siegel’s definition of mind, which challenges this conventional view as being too limited.

Siegel argues that viewing the mind as arising solely from the activity of the brain fails to do justice to the mind’s complexity. While the brain is very likely the physical structure that registers conscious self-awareness, the mind consists of input from other sources as well. Siegel defines mind as the “flow of information and energy” among our brains, our bodies, and one another—that is, our social interactions with other people. In other words, the mind and brain aren’t two sides of a coin. The mind arises from brain (neural) activity, to be sure, but the mind also consists of input from the body and from our relationships—from the meeting of minds, you might say.

This argument makes sense for several reasons. First, while our bodies connect to our brains though neurons that send information (such as commands) from the brain to the body, and which deliver information from the body to the brain, the body can act independently of the brain to some degree. This is especially true of the enteric system, the digestive system at the core of our bodies, which includes the esophagus, stomach, and intestines (large and small), among other structures.

The enteric system actually resembles the central nervous system (brain and spinal cord), containing neurons that are more brain-like than those anywhere else in body, and that connect with one another to form a system relatively independent of the brain. Every neurochemical found in the brain is also found in the digestive system; in fact, most of our serotonin, the neurotransmitter thought to cause depression if there are insufficient quantities available, is found in digestive areas. An anxious girl at the Meryton ball who felt butterflies in her stomach would have demonstrated the enteric system informing the brain about her nervousness rather than the other way around. Psychological reactions in the stomach and digestive system might depend very minimally on brain activity.

The enteric system might be a particularly brain-like bodily area, but many areas of our bodies participate in thinking. Recall that emotions begin as physiological responses that we subsequently register as feelings. Even before Siegel redefined mind, many mind-brain scientists accepted the phenomenon of embodied cognition, the belief that thought and behavior depend on bodily responses to a particular environment as well as neural activity.

That our social interactions belong to the definition of mind also makes sense. Our minds and our brains depend on social contact for both development and well-being. If you confine an infant in a room, providing food alone with no nurture or human contact, you’ll irrevocably damage his brain within a very short time. In fact, it’s unlikely he’d survive, and if he did, he’d be severely impaired, compromised for life. Social contact continues to be vitally influential throughout our lives, influencing feeling, mood, perception, and cognition. We’re designed to pay attention to one another—it’s an inherent part of what constitutes our minds, both developmentally and moment by moment.

Psychiatrist Lewis Mehl-Madrona, while not specifically defining mind, supports Siegel’s definition with an analogy: “People are neurons in a social brain.” By viewing activity within this social brain as a building block of individual minds, Siegel takes this insight to the next level. Activity within the social brain and activity within individual mind-brains continually constitute one another in an ongoing feedback loop, a process called “recursivity.” (As you’ll see in a moment, recursivity is a defining feature of the mind-brain.) There’s no mind without other minds; John Donne’s famous line “No man is an island” applies to the neurological and psychological findings of our own day. I think Austen would have approved of Siegel’s theory, knowing as she does that the real stuff of our lives can be found in our relationships with one another.

Fundamentals about the brain are nevertheless helpful to understanding the mind, even if mind and brain are different entities. You don’t need to remember everything in this chapter about the brain, but reading through will give you the basics of how the brain functions. So now, to return to Darcy’s contemptuous glance.

For Darcy to see Elizabeth, sensory information from the outside world must reach the conscious parts of his brain. This means that when Darcy “looked for a moment” at Elizabeth, light traveled through the lens of his eye and hit its back surface, the retina. The deepest layer of the retina (nearest the brain) contains millions of photoreceptor cells, some of which are sensitive to light and some to darkness. These cells, and all the others involved in the brain’s processing, are called neurons, or nerve cells. (There are other kinds of brain cells, glial cells, involved in maintenance, but neurons appear to be involved in thought, affect, perception, and action.)

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Elizabeth’s image caused these photoreceptor neurons to fire, the conventional term for referring to the activation of neurons, creating the pattern of her image, much like a photo negative. This in turn activated the retina’s output neurons (called ganglions), which sent the image to areas deep inside the brain via the optic nerve, which isn’t really a single nerve but a collection of neurons. The impulses carrying Elizabeth’s image then traveled upward through many layers of the brain to reach the visual cortex, the brain’s highest and most advanced visual processing area.

The journey still wasn’t over. The visual cortex as a whole contains a topographic map of the visual field, which means that the neurons in a given area correspond to specific areas in the visual field; each cell is responsible for a small part of the picture, like the pieces of a jigsaw puzzle. Cells of the visual cortex are also specialized to process different aspects of perception, such as form, color, or movement. For instance, the primary visual cortex, known as V1, analyzes patterns and the orientation of forms, such as whether Elizabeth is facing right or left, and whether the lace she’s wearing has a regular design. Other specialized cortical areas are needed for recognition of objects and faces, and for the perception of three-dimensional forms. Only when all the different parts of the image contained in various electrical nerve impulses were assembled in jigsaw-puzzle style was Darcy able to see Elizabeth, “not handsome enough to tempt me.”

In short, Darcy didn’t perceive an image of Elizabeth wholesale, like a boa constrictor eats its prey, but rather, patterns of light and darkness that struck his retina were turned into electrical impulses that, after being registered and assembled by different visual centers in his brain, eventually formed the image of Elizabeth. Yet Darcy remained oblivious to the complicated workings of visual processing. And so it is with most of our perceptions. The workings of many brain areas are involved, yet we experience an end result without being aware of the steps needed to produce it.

Take another example of a brain process: memory. We don’t record memories like a video recorder and then play them back; rather, we fashion prior perceptions and experiences anew each time we think of them. The constructed—not recorded—nature of memory leaves plenty of opportunity for error, omission, and variation: You can be sure that if you asked Darcy about the Meryton ball, his account would differ significantly from Bingley’s. And when you remember an event, the same areas that were active in registering your initial perceptions work to re-create it. When Darcy later recalls Elizabeth’s fine eyes, the visual centers that sprang into action when he first noticed how pretty they are reactivate to form the image in his brain, although in a modified, less intense form. The “mind’s eye,” an expression that predates the field of neuroscience, is accurate. The mind really does have an eye insofar as the brain has visual processing centers that create mental images. These areas are active whether what we see is real, remembered, or imaginary.

One more crucial point to remember about the mind-brain: There’s nothing inside the skull but the brain itself, no little person in a control tower, no vital spirits or other immaterial substances, no ghost in the machine, just a brain gathering information and sending information. Yet the absence of an animating spirit in the conventional sense takes away none of the wonder. Rather than simply glorying in the talents of the human mind-brain, as we’ve done since time immemorial, we now ask how its abilities can arise from the same everyday processes seen in other mammals, whose mind-brains comprise the same elements as our own: brains, bodies, and social interactions. A group of chimps can’t write Hamlet, or Pride and Prejudice, no matter how much time they’re given in front of a computer; you need a human mind for that (and an extraordinary one to boot).§ But despite the commonplace materials our minds are made of, even minds like Shakespeare’s and Austen’s, we can still say with Hamlet, “What a piece of work is man” (and woman!).

THE DANCE OF NEURONS

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Scientists tend to speak of brain areas as if they were factories where workers accomplish different jobs, but actually, these areas are made of neurons, and neurons alone get the work done. We can’t tell the dancer from the dance, neurons from the brain, and it’s the endless dance of neural activity that’s vital to making us who we are. Similarly, when scientists talk about information being stored in neural circuits, they mean that a particular thought, feeling, or action will happen if a particular configuration of neurons activates.

Neurons are the information processing cells of the brain and nervous system; through electrical impulses, they both give and receive information that travels from one neuron to the next, or from a neuron to an area of the body. You might compare this to the sounds of our voices traveling along telephone wires (or wireless systems), which are transformed into energy that’s reconstituted as information at the receiving end. And so knowledge about whether or not Elizabeth is pretty (knowledge involving the entire mind) travels from Elizabeth herself (social interaction, a glance at a ball) to photoreceptor neurons in the eye (body), to cortical neurons that assemble her image (brain). There’s an average of 86 billion neurons in the human brain.

Neurons come in many different shapes and sizes, but most of the neurons in vertebrate brains are multipolar. They consist of a cell body, which contains a nucleus; dendrites, which are small extensions that receive electrical impulses through receptors; and an axon, a long, tube-like structure with extensions for sending electrical impulses on to other neurons or body areas. Axons come in all different lengths, and some are long enough to conduct impulses from the brain to the far reaches of the limbs. The multipolar neuron is probably what you think of if you think of neurons at all; its familiarity is shown by its incarnation as a stuffed toy sold by the Giant Microbes company. (They also sell blood cells, bone cells, E. coli, and other cells that nonscientists are likely to recognize from the popular press.)

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Every one of your thoughts, feelings, and actions involves a cycle of neural activity that begins with a stimulus of some kind, such as exchanging a glance with a young lady at a ball or remembering her fine eyes. This causes electrical current to flow into a neuron via the activity of chemicals called neurotransmitters; these can be likened to other materials that conduct electricity, such as metal or water. For an electrical impulse to travel from neuron to neuron, it must cross a tiny space called a synapse. Small amounts of neurotransmitter are secreted from pouches called vesicles, located in the axons of a sending, or presynaptic, neuron and received by receptors in the dendrites or the cell body of a receiving (postsynaptic) neuron. But the transfer of energy from neuron to neuron will take place only if the postsynaptic neuron has receptors for that particular neurotransmitter. Neurotransmitters are like keys, and receptors are like locks; the gates will open only if lock and key fit together.

Most neurons connect to other nearby neurons within a specific area of the brain rather than connecting directly to neurons that have contact with the outside world. There are input neurons that take in information from the environment, as in vision, and neurons that generate information from within, as in memory. These neurons connect to other intermediary neurons, called interneurons, and interneurons connect to output neurons that implement a response.

Interneurons often add to the information they receive before sending it on. Darcy’s initial visual intake sent information through many layers of interneurons in various brain areas before reaching the output neurons in the visual cortex that assembled Elizabeth’s image. However, some neurons connect directly to target areas, such as the nerve cell that generates the knee-jerk reflex, tested at your yearly physical. But most human brain activity involves interneuron activity for both receiving and sending information before the final output.

The majority of neurons are specialized in terms of functions they enable; as we saw, the photoreceptor cells that sent the electrical impulses that led to Darcy’s seeing Elizabeth began their activity in response to light, an environmental stimulus. Other kinds of neurons deal with more abstract psychological processes. But whether we’re looking at someone’s fine eyes, remembering a humiliating faux pas, worrying about the future, or dancing at a ball, neurons are part of the process, as they calculate input in order to generate responses, moment by moment.

Neural signals can inhibit as well as stimulate the activity of other neurons. In other words, the electrical signal can convey the message “Don’t fire” as well as the message “Fire.” A receiving neuron’s activity, or lack of activity, results from the activity of all the neurotransmitters that are converging on the cell at the same time. That is to say, in each neuron, receiving impulses are literally added together, with inhibitory impulses (Don’t fire!) subtracting from the total of excitatory ones (Fire!). This means that neural activity is the result of each neuron’s calculation of the excitatory and inhibitory signals it receives; a neuron will fire if the sum of all of its activating impulses is strong enough. Each neuron receives and sends thousands of impulses in any given moment. The math involved in neural activity is one reason many cognitive scientists believe the brain is computational, that it functions like a computer.

You might think that criticism of a physiological, computational view of the brain would come from humanists and literary people. The thought of Darcy and Elizabeth as walking computers, calculating the sum of neural activity, is likely to irritate many Austen enthusiasts. But the critique can be found within science itself, from the undeniable observation that in the brain, the sum of neural activity is greater than its parts. The same is true of the mind, which includes the body and social interaction in addition to brain activity.

Take what neuroscientists call “the hard problem of consciousness,” how awareness emerges from the workings of the mind-brain. Even if we were able to trace the activity of every one of the brain’s billions of neurons and, incorporating Siegel’s views, include the input we receive from our bodies and social information in the environment, we still couldn’t say how our awareness of our perceptions (called qualia) and of ourselves emerges from the interaction of these elements. Even the best theoretical explanations can’t account for how this happens, which is why figuring out how consciousness emerges is called a “hard” problem. And this is an understatement, because even if you could trace the activity of every neuron in the brain and the body, in addition to calculating every factor in the environment (and of course these tasks are impossible), you still couldn’t explain how such activity translates into “how things feel”—the phenomenology of experience. The gap between the workings of the mind-brain and the quality of our experiences might be one that we’ll never bridge.

DYNAMIC DARCY

Those who fail to appreciate Austen foolishly (in my view) assume that her novels are dull because her situations tend to be relatively ordinary: Darcy moves into the neighborhood; Anne Elliot’s family is forced to rent the family home in order to pay their debts (Persuasion); Catherine Morland, who’s never traveled, has the opportunity to visit Bath (Northanger Abbey). The action is primarily psychological; Austen focuses on feelings and insights that occur as a result of social interactions in a relatively limited milieu. But for humans and other primates, this is the stuff of our most important events. Austen knew that the real drama of our lives most often comes from our everyday encounters with other people.

Such encounters bring challenges in their wake. And people, including fictional people, have various and innovative ways of responding. If this were not the case, every novel would be more or less the same—indeed, we’d have no novels, for what would be the point? We read because we’re interested in people and the choices they make as they attempt to navigate their way to happiness.

Psychology explains many of the factors that influence our responses, but for the moment, let’s remain at the nuts-and-bolts level of brain function. At this physiological level, we can say that people have a wide range of possible responses to experience because the brain is what is known as a “complex system,” also called a “dynamic system,” a concept that originated in the field of thermodynamics. (The study of such systems is known as general systems theory.) The mind is also a complex system, but we’ll focus on the brain to begin with.

Complex systems are self-organizing systems. As I mentioned earlier, there’s no little person, no “mini-me” inside Darcy’s skull directing the activities of his brain: And yet despite the absence of this cognitive “author,” Darcy and all of us with healthy brains function incredibly well, at least from a mechanical if not always from a social or psychological perspective.

Heating a pot of water provides a nonorganic example of a complex system that anyone who’s waited for pasta water to boil will be familiar with. Without heat, the water molecules move in random ways. But when the water is heated, it forms small bubbles, then large bubbles, then agitated coils of water. No one is shaping the water in this way, but it forms noticeable patterns nevertheless. And the sequence repeats every time you boil a pot of water.

Of course, the activity of a complex system is governed by the restraints that are built in to the materials it’s made of. Water in a pot is never going to read Austen, even if it’s governed by the same principles as the brain. In the case of brains, restraints include basic anatomy and an individual’s genetic heritage, the latter estimated to have an equal influence with the environment in the majority of instances.

In complex systems, including both boiling water and the brain, self-organization occurs as the result of feedback loops that affect the functioning of the entire system, a phenomenon called “recursivity.” A neuron that sends a signal out, either to other neurons or to an area in the body such as a gland or a muscle, is simultaneously taking signals in from other neurons, including those that bring information about the environment or the body or other parts of the brain. As signals continue to circulate, some of the impulses received will be responses that have themselves been influenced by the neuron’s own output. As noted, this will almost always happen indirectly through relays of interneurons.

Neural networks are so dense and complicated that any one brain area is constantly sending and receiving information to and from a multitude of other areas. So when a complex system confronts a stimulus, as the brain does each and every moment, the changes induced don’t ripple through the system one after another like dominos falling; this is linearity. Rather, recursivity, self-organization through feedback loops, means that the stimulus affects the system as a whole. The result is an endless circuit of influence and adjustment, happening at lightning speed.

The mind is also a complex system. To add the mind back in to the equation, we just need to consider bodily and social activity as part of the endless feedback loop that makes us who we are each and every moment. If you could have a freeze-frame photo of all the billions of connections that are being made within and between brains, bodies, and people, you’d have a “screenshot of the self,” of the configuration of inputs that forms our identity—all of our perceptions, intentions, thoughts, and feelings (including emotions and bodily sensations), at that particular moment. But of course, within milliseconds, the picture will have changed.

When life confronts us with experiences that can’t be absorbed in a routine manner by smooth and continuous reorganization of a complex system, which for the mind translates into taking things in stride and coping well in terms of thought and behavior, we can respond in one of the three ways available to all complex systems: with rigidity, chaos, or adaptive reorganization. Rigidity means failing to change in ways that accommodate or even acknowledge the challenge—rigidly maintaining old patterns. When Elizabeth rejects her foolish cousin Mr. Collins’s proposal of marriage, he hunkers down and clings to what he knows. He’s so locked in to certain modes of perception that he literally can’t absorb what she tells him—that she doesn’t want to be his wife.

By contrast, in Sense and Sensibility, when Willoughby rejects Marianne in a very cruel and public way by snubbing her at a ball, Marianne has a chaotic reaction, which means that she can’t assimilate the information and move on. She’s completely overwhelmed, and she ceases to function well or normally. Like Mr. Collins, she clings to cherished beliefs to begin with, thinking that someone must have maligned her to Willoughby, and that’s why he’s rejecting her. Never mind that such behavior on his part would still be wrong, trauma doesn’t stop to examine nuances. Once Marianne realizes that Willoughby alone is responsible for his behavior, she descends into a deep depression. She’s lost in the chaos of her pain, which takes her over entirely.

Marianne’s state of mind paradoxically involves rigidity as well as chaos, the lack of ability to function normally, in that it follows predictable patterns—sad mood, lack of responsiveness, loss of the capacity for pleasure (anhedonia), loss of appetite. These symptoms indicate a deficit in her powers of emotional regulation, an absence of organization, that leads to the rigidity of depression. Such maladaptive expressions of system overload can be characterized as both chaotic and rigid.

Like Mr. Collins and Marianne, Darcy is also rejected after a proposal (to Elizabeth). But in his case, the rejection initiates a massive reorganization of thought, perception, emotion, and behavior that leads to a better system and a better man. Although Darcy is initially completely taken aback and furious at the rejection, he thinks long and hard about what’s happened, and he changes as a result. Elizabeth immediately notices how different he is when they later encounter one another at Pemberley, Darcy’s estate. She’s surprised by his gracious and welcoming attitude, especially toward her aunt and uncle, the Gardiners, despite Mr. Gardiner’s being “in trade” and so beneath Darcy in terms of social status. Darcy has shed his defensive, protective armor and is much more open to experience. He takes in information and decides on appropriate responses rather than automatically distancing himself and shutting others out.

After Elizabeth’s rejection, Darcy might well have chugged along, gotten over it, and asked another woman to be his wife à la Mr. Collins—perhaps Miss Bingley’s dreams would have come true after all. Or he might have become deeply depressed and dysfunctional, like Marianne after Willoughby’s rebuff. Instead, he becomes a kinder and happier person by processing his experience in ways that enable him to substantially change for the better. Such change indicates flexibility of mind, and it augurs well for his ability to respond in productive ways to experience in the future.

In case we’re tempted to attribute Darcy’s transformation to an effort to capture Elizabeth’s heart, a blip that will have no lasting effects, Austen makes sure we know that Darcy has processed his experiences—rethought his attitudes and behavior—and changed as a result. He tells Elizabeth,

As a child I was taught what was right, but I was not taught to correct my temper. I was given good principles, but left to follow them in pride and conceit. . . I was spoilt by my parents, who . . . allowed, encouraged, almost taught me to be selfish and overbearing, to care for none beyond my own family circle, to think meanly of all the rest of the world, to wish at least to think meanly of their sense and worth compared with my own. Such I was, from eight to eight and twenty; and such I might still have been but for you, dearest, loveliest Elizabeth! What do I not owe you!

Darcy’s mind reorganizes itself to accommodate his experience productively. And he can develop with such leaps and bounds precisely because he—or rather his mind—is a complex system.

When you think about it, one rejection in the context of a lifetime of experiences shouldn’t logically be a life-changing event. And for Mr. Collins, it isn’t. But obviously in Darcy’s case, the rejection ripples through his mind touching on all sorts of issues, perceptions, memories, feelings, desires, and so forth, eventually altering him in dramatic ways. The change is system-wide and pervasive.

Darcy’s transformation points to one of the remarkable qualities of complex systems, particularly complicated ones like the mind and brain: Seemingly small events can trigger huge changes. We see this with other Austen characters as well; they learn, they grow, they become their best selves, and these changes appear to be disproportionately large compared to the events that instigate them. The recursivity of the mind, its operation through feedback loops, makes this possible. In Darcy’s case, reorganization is for the better. But recursivity also means that a single negative or traumatic event can have a seemingly disproportionate negative effect, as with a traumatic experience.

Change can also be gradual and cumulative, another quality of complex systems that’s useful to understanding human psychology. Small changes can occur on an ongoing basis until they form a critical mass, and then we notice that the system has substantially progressed from an earlier point. Darcy falls for Elizabeth, hard and fast. But Elizabeth’s love develops slowly, as she gradually realizes Darcy’s worth and their compatibility as a couple, until her thoughts reach a tipping point, a moment at which she knows that Darcy is the man she wants to marry.

ALL TOGETHER NOW: NEURAL INTEGRATION

Darcy’s response to the challenge of rejection is a better one than that of Mr. Collins or Marianne because it’s a more complex one. I mean this in both a colloquial and a technical sense. Both Mr. Collins and Marianne respond to crisis in ways that are simple because they’re straightforward and predictable, seen in countless other people, real and fictional. Mr. Collins can’t adjust his perceptions or expectations because his character is inflexible, while Marianne’s despair follows well-known and repetitive patterns that have been seen in countless people who become depressed. This predictability has enabled psychologists to come up with diagnostic criteria for classifying depression as a disorder.

But Darcy’s response is more complex because it doesn’t follow an expected or familiar pattern. As a result of the shock of rejection, he evolves new ways of responding to people and events, which involve multiple and variable adjustments in thoughts, feelings, values, and behavior. These differ from his habitual reactions, as well as from well-known categorizable, if chaotic, conditions such as depression.

The terminology here is confusing. Of course, both Marianne and Mr. Collins have complex mind-brains in the sense that they have human brains that function through neural feedback loops—recursivity—and so their brains are, by definition, complex. In addition to more localized circuits among brain areas, energy and information are constantly flowing bidirectionally through every brain between bottom and top, left and right (between the hemispheres), and front and back; we wouldn’t function at all without such neural activity.

But Darcy’s mind-brain exhibits greater and more successful complexity than those of his counterparts. Success in a complex system refers to the extent to which it expands in the best way possible, which involves developing differentiated, specialized modules and greater interaction among those areas. When water boils, it begins to function as a whole (interaction) rather than as the collection of separate bubbles we see when at the beginning of the process, and it forms noticeable and various patterns (differentiation) through the eruptions of coils of water. All boiling water will function on par, but some brains will specialize and interconnect (and the latter is particularly important) more successfully than others. Such interconnection is known as neural integration.

Darcy’s mind demonstrates complexity as defined in this scientific sense, reconfiguring and creating new neural connections in the body and brain, which means new ways of thinking, feeling, and relating. The mind-brain’s recursivity means that these new modes of being further affect social interaction, brain, body, and behavior—and so the cycle continues. Darcy’s responses to experience become more differentiated from familiar or default patterns and integrated because the brain and body and sociality work together to further his well-being. In terms of the brain alone, a complex response to experience calls on concerted activity from a greater number of densely interconnected brain areas that work together than we see with rigid or chaotic reactions.

Complexity gives Darcy the ability to think creatively and respond in productive and appropriate ways to his experiences. Greater complexity translates into the flexible cognitive and emotional flexibility as well as the successful regulation of emotion. By contrast, both Mr. Collins and Marianne deal with their crises with default modes, rigidity and chaos (depression), one-size-fits-all reactions applied across a wide variety of experiences. Darcy himself emerges from a Collins-like rigidity. You can see this if you compare the automatic, almost rote nature of his responses at the Meryton ball with the quick, varied, and appropriate interactions he has with Elizabeth and the Gardiners when they encounter him at Pemberley.

Greater complexity generally translates into greater stability and flexibility, which means better functioning. As Darcy’s mind-brain moves toward greater complexity, it calls upon more working parts, if you will, parts that coordinate their activity in diverse neurological and psychological domains. At the Meryton ball, Darcy is on automatic pilot, responding to experience without really thinking about his words or his actions, guided by feelings that he hasn’t really considered. At Pemberley, he behaves with thoughtfulness and consideration; cognitive and emotional brain areas are fully interconnected and working together in concert with bodily responses and social awareness.

The evolution of the human brain offers an illustration of complexity that’s easy to grasp because it shows a clear connection between anatomy and function, one that we don’t have the tools to trace in terms of the functioning of mind as a whole. Primate brains evolved by developing neural areas that became progressively more specialized and differentiated as different species evolved. The ability to produce speech stems from a part of the human brain known as Broca’s area; we find a corresponding area, also responsible for communication in chimpanzees and other great apes. These cousins of ours communicate with one another through vocalizations, social signals, and gestures. With training, chimps in captivity can even be taught to use American Sign Language. (But they can’t use language in the way humans can; they can learn vocabulary and convey ideas with words using sign language, but they don’t have syntax, the knowledge of grammatical rules for forming sentences). Chimpanzees tend to use communicative gestures with their right hands, which demonstrates left-brain dominance for communication, just as is found in humans. They also communicate meaningfully, just like humans, and indeed their gestures have consistent meanings, like the words in a language. All of this means that chimps have a language-ready brain, which began to evolve in the branch of our common ancestor that gradually became human.

The fact that an area corresponding to Broca’s area orchestrates communication in chimps shows that evolution is thrifty, reusing and recycling materials that are already there. In the course of evolution, brains didn’t as a rule develop complexity by generating additional brain areas, but by adding neurons to and, even more important, increasing interconnections within and between existing areas. The superior and specialized functioning of Broca’s area in humans, the “complexity” of our capacity for language, results from a greater number of neurons and increased connectivity, denser and more complicated wiring.

Density also applies to developing increased complexity within individual brains. Although neurogenesis, the creation of new neurons, takes place in brains, especially during childhood, complexity tends more often to develop through greater interconnection among existing neurons. Darcy’s brain functions better, with greater complexity, because it makes a greater number of neural connections than does Mr. Collins’s brain. As its neural wiring becomes denser, Darcy’s repertory of available responses correspondingly increases. Darcy’s neural circuitry builds more roads within the brain, and so he has more routes available on his journey (more neural circuitry) and more towns that can be visited. This means a greater number of options for response, which confers flexibility as well as resiliency to setbacks and rejections. If one road leads to a dead end, take another.

Very simple behaviors such as the knee-jerk reflex involve only one neural pathway running from sensory input neurons in the knee to motor output neurons in the brain. But much of the brain’s processing, which includes taking in information and generating responses, happens through processing that takes place simultaneously in different areas and is then assembled. This is called parallel distributed processing.

The pooling of information from different areas of the brain is known as neural integration, a feature found in all vertebrates, creatures that possess a central nervous system: a brain and a spinal cord. For the brain to coordinate input from many different areas, there needs to be an efficient way of bringing neural signals together. To return to the roads metaphor, if you lived in a small rural town in the south of eighteenth-century England, you couldn’t travel a direct route to Gretna Green in Scotland, where eloping couples could be hastily and legally married, because the roads for such a journey didn’t exist. So you’d first have to travel to the country’s hub, London, and from there find roads leading to Scotland. That’s why when Mr. Bennet searches for Lydia and Wickham (they’ve eloped), he travels to London where he knows they’ll first have to stop. In the same way, neural signals from different areas of the brain, which have been processed in parallel, need to travel to hubs, known as association areas, where information is assembled and sent along for further processing. But association is almost never the sole function of these areas—far from it. In the process of putting information together, they add their input.

Neural integration is good for the brain, and so for the whole person—the more of it there is, the better. As Siegel explains, the more interconnected the brain is, the more that various circuits work together, the greater the flow of energy and information will be throughout, and the better it will function. The reverse is true as well: Both pathological conditions and less than optimal responses to ordinary experience are often characterized by insufficient neural integration. Think of a quadrille, a popular eighteenth-century dance for four couples. If even only one dancer misses a cue, he’ll compromise the dance for everyone. Similarly, if there’s a brain area or circuit that isn’t working in concert with other parts of the system in the way it should be, this can compromise the functioning of the whole.

You’re likely familiar with a very well-known instance of the failure of neural integration: post-traumatic stress disorder. For the unfortunate sufferers of this condition, traumatic memories erupt in flashbacks and dreams because the original traumatic experience can’t be processed or “taken in” by various parts of the brain, particularly higher cortical areas. Traumatic experience is registered in emotional and bodily memory, subcortical systems that have no sense of time. And so the sufferer experiences disturbing, intrusive symptoms such as panic attacks and flashbacks that make the experience seem part of the present rather than the past. Neural integration is one route to healing; if cognitive areas can process traumatic experience, for instance, by putting feelings into words that are part of a narrative with a beginning, middle, and end, symptoms are often ameliorated. The experience then becomes part of the past rather than the present.

Marianne’s depression similarly involves the shutdown of cognitive areas; negative emotions take over her mind at the expense of all other functions. Other less extreme failures of neural integration happen routinely, as in the psychological process of denial, a defense mechanism that means failing to notice something because it is threatening or disturbing. Mr. Collins must compromise perception and logic to turn Elizabeth’s “no” into “yes,” even if young ladies did often refuse proposals they eventually meant to accept, as he believes Elizabeth is doing. The affect accompanying her refusal, conveyed by social signals such as facial expression and body language as well as speech, should have convinced him that she was sincere.

Neural integration and parallel distributed processing (which together generate the simultaneous processing and subsequent pooling of information) refer to mind as well as brain. Darcy’s glance at Elizabeth, which is itself processed in parallel and assembled in the brain, is only one component of his behavior at that moment. At the same time, he’s conscious of his resentment for having to attend the ball and evaluating Elizabeth’s looks. Had he simply looked at Elizabeth and seen a young lady sitting there, he wouldn’t have made a nasty comment. But vision is integrated with information from other mind-brain areas, such as the emotional areas that tell Darcy he’d rather be elsewhere, from body areas necessary for perception and emotion, and from the social context of viewing a young lady at a ball while knowing that he’s the door prize.

Of course, not everyone who responds with complexity to experience, who has an optimal flow of energy and information throughout the mind-brain, is going to dramatically improve their character or marry the partner of their dreams. Jumps between levels of analysis should be handled with care. But in general, complexity of thought in both the colloquial and technical senses leads to more flexibility and more numerous ways of responding, characteristics that are crucial elements of social intelligence.

HEBB’S RULE

Pride and Prejudice has been characterized as a novel of manners, a marriage plot novel, a romantic (and Romantic-era) novel, a didactic novel, a work of great literature, and probably in other ways as well. I’ll add one more to the list: Pride and Prejudice can be characterized as a Hebbian novel, named for one of the founding fathers of neuroscience, Donald Hebb (1904–1985).

Dynamic systems move toward complexity through change, growth, and development. But they are characterized by continuity as well as flux. A foundational principle of neuroscience, known as Hebb’s rule, accounts for the continuity of patterns of thought, feeling, and behavior that characterize the mind-brain. Donald Hebb’s important discovery is widely known in the form of a catchy saying: “Neurons that fire together wire together.” This means that once a group of neurons has activated at the same time, they’re more likely to activate together in the future. Habitual patterns of neural activation are called attractor states.

Continuously recurring neural patterns generate the enduring traits that make up personality. Extraversion, introversion, optimism, pessimism, to name a few features, exist because certain neural patterns tend to repeat. Recurring neural patterns also enable learning; knowledge, like memory, depends on the activation of neural patterns that have previously fired. You repeat certain neural patterns every time you recall a basic fact such as London is the capital city of England. This principle applies in far more complex forms of learning, as well as for habitual ways of perceiving, feeling, and behaving. In short, attractor states generate much of our minds, who we are and what we know. They enable constancy and stability in the mind-brain; they enable us to have a consistent sense of our own identity even though our mind-brains are in a continuous state of flux.

Attractor states are of course responsible for the stubbornness of prejudice. The word prejudice as we use it today usually refers to an opinion formed without information. In Austen’s day an alternative definition, according to the Oxford English Dictionary, was “a premature or hasty judgment.” This aptly describes Elizabeth’s behavior. She forms a devastating and persistent opinion of Darcy on their first meeting at the Meryton ball, and this is reinforced every time she meets him or thinks about him. Neural pathways that would allow her to correct her first impression can’t compete with entrenched responses. Hebb’s rule trounces incoming information.

However, this is only part of the story. Although at the neurological level, Hebb’s rule explains the persistence of misguided thoughts and feelings—Elizabeth has an attractor state for seeing Darcy as negatively as possible—this explanation is insufficient on its own to account for Elizabeth’s prejudice. Whether as a reader of Austen or an observer of human nature (and if you’re reading this, you’re bound to fit at least one of these categories), you’ve surely noticed that some misconceptions can be easily corrected while others are ingrained. Certainly, Elizabeth’s mind-brain predisposed her to think poorly of Darcy after their first meeting. But she soon had access to additional information that should have contradicted her view of him as past redemption. She herself eventually realizes that she shouldn’t have trusted Wickham’s story. Elizabeth’s misconceptions become entrenched from the start.

This is because they serve a purpose. When Elizabeth complains to her friend Charlotte about Darcy’s excessive pride, Charlotte points out that given his money, rank, and personal attractiveness, Darcy “has a right to be proud.” With her characteristic quickness and wit, Elizabeth rejoins, “That is very true, . . . and I could easily forgive his pride, if he had not mortified mine.” The word mortified is telling; it comes from the Latin meaning “to kill,” which points to the damaging nature of Darcy’s remark. Mortification doesn’t kill, but it’s very painful, a form of humiliation much more wounding than simple embarrassment. Darcy’s words didn’t kill Elizabeth, but they nevertheless annihilated her symbolically, pegging her as worthless.

And so Elizabeth misses vital information that would contradict her first impression because she’s invested in thinking of Darcy as a bad man. Her “prejudice” is a form of self-protection. If she can believe that he’s bad, then she can believe that his judgment is faulty, especially his judgment of her. Elizabeth’s prejudice is also a form of revenge; if Darcy thinks ill of her, she’ll return the favor. But as Elizabeth realizes, it’s unusual for her to misconstrue persons and situations so wrongly; in the usual course of events, she “prides” herself on her “discernment.”

We can’t yet explain how such filtering mechanisms work, exactly how perceptions are distorted. But we know, as in Elizabeth’s case, that feelings are responsible. Emotions drive not only the social brain, but the thinking brain as well. Fortunately, emotions drive change and growth as well as distortion and prejudice, as we see in the story that follows that first unfortunate glance at the Meryton ball.

We Janeites accept that we thrill to the romance of seeing a deserving heroine enjoy her Cinderella-esque ending. And many of us additionally congratulate Elizabeth for winning her happy ending on the basis of character and intellect rather than feminine looks or wiles—her eyes owe much of their beauty to her liveliness of mind. But there’s an equally thrilling, if more muted story being told about the power of love and, broadly speaking, all positive relationships, to transform us in extraordinary, unexpected ways.

FINDING YOUR WAY AROUND THE BRAIN (AN INTERLUDE)

Understanding neurons, complex systems, and Hebb’s rule provides crucial information about how the brain functions. A basic knowledge of brain anatomy can be additionally helpful to understanding the neural processes that contribute to making us who we are. However, those readers who’d rather not pursue this topic can move on to the next chapter; brain areas will be explained as we encounter them.

The brain divides roughly into three areas, which developed chronologically in the course of evolution. They go from lower to higher, and inner to outer; in general, the higher and more exterior these areas, the more advanced the functions they oversee. We can imagine the spatial relationship of these areas to one another by visualizing a cauliflower. (Actually, the brain looks most like a walnut, a food that’s good for talking about the hemispheres [see below]. But a walnut fails to portray the brain’s levels correctly!)

The first part of the brain to develop in evolutionary time (phylogenetically) was the brainstem (the stalk of the cauliflower), which is responsible for a variety of survival behaviors, such as breathing, sleep, arousal, and the regulation of heartbeat. The brainstem also participates in bringing information about the world to other parts of the brain and conveying commands from these higher levels to the body; it’s an information superhighway, continuous with the spinal cord, whose neurons reach throughout our bodies. Brainstem activity is involuntary and subconscious. Reptile brains are mainly brainstem, with very primitive versions of other structures. So if you think your pet snake loves you, you’re projecting your own feelings onto a creature who’s cold-blooded in emotional as well as physiological terms. Unlike mammals, reptiles lack the neural machinery needed to form emotional bonds. (However, I have a friend who swears that bearded dragons relate and cuddle in mammalian ways. Perhaps mammalian hubris is as bad as anthropocentrism!)

Next, moving upward, are the subcortical areas, the core of the cauliflower, which includes the structures that make up the limbic system; these areas are key to the processing of emotion. They include the hypothalamus, which activates the release of important neurochemicals and hormones, such as the hormone adrenaline. This chemical messenger is involved in producing excitatory responses of both positive and negative kinds—joy as well as fear. The hippocampus forms new episodic memories, memories of people and events. And the amygdala, one of our two “star players” (see below) is central in the activation and recognition of fear as well as other excitatory responses. The amygdala tells the hypothalamus to send the message to release adrenaline. It’s also one of the brain structures that tags events as important to remember.

The development of limbic brain areas in mammals—this area used to be known as the mammalian brain—means that mammals share many of the same basic emotions as humans, as the study of animal behavior and neuroscience have suggested. This accounts for the sociality of mammals, as seen especially in their nurturing of offspring and their bonding with one another and with humans. Unlike your pet snake, your dog really does love you (and don’t believe those naysayer skeptical scientists who continually underestimate animal capabilities). Perhaps this exonerates some of Austen’s sporting types such as Charles Musgrove in Persuasion, who’d rather spend time with his dogs than with his self-centered and tedious wife. The experience might indeed be more emotionally gratifying. The limbic areas are certainly not the only brain areas dedicated to the processing of emotions, but they’re important subcortical ones.

Finally, the cerebral cortex, made up of the flowers of the cauliflower, is the most advanced and recently developed of brain areas (it too has raised and recessed areas, even though it isn’t as good a brain look-alike as the walnut). It’s the outermost layer of the brain, and it wraps around subcortical areas like a blanket. It’s often called the neocortex in humans because it is so new (neo) and advanced compared to the cortex in other vertebrates. The cortex exists in reptiles as a single-cell layer toward the back of the brain. In most mammals, it’s a thin layer of tissue surrounding the limbic structures. But in humans and other primates, it’s huge in relation to other brain areas.

The cortex expanded in “advanced” mammals by folding in on itself to form grooves (sulci) and raised areas (gyri), thereby providing more surface area and a greater number of cells, resulting in more brainpower. This gives the wrinkly, convoluted appearance most of us picture when we think of the brain. If you compare a hamster brain to a dog brain, you’ll see that the rodent has fewer of these convolutions. The more developed the cerebral cortex, the greater the number of sulci and gyri, and the greater the number of capabilities, including social capabilities, an animal possesses. Think of the difference between a pet hamster and the family dog. One has a more developed cerebral cortex, the other, only a very basic one.

Social capability refers not only to depth of feeling, although this is often the case, but to the multifaceted ways that we interact with one another. For despite the common substrate of sociality and emotion we share with other mammals, human cognition adds options for response not found in other species. Perhaps relationships among some of these other animals have dimensions of which humans are unaware, and which surpass ours in complexity and sophistication—who’s to say? Although our anthropocentric species has underestimated the mind-brains of other species for most of our time on earth (and many continue to do so), I think that even those who appreciate how smart (cognitively and emotionally) many other species of animals are—and I consider myself in this company—would still grant that the complexities of feeling found in Pride and Prejudice are likely not found in other species, not even in chimpanzees and bonobos, who share almost ninety-nine percent of DNA with humans. We can be relatively certain that no chimpanzee will ever type out Pride and Prejudice not only because chimps lack the language and intelligence to do so, but just as important, because as far as we can tell, they lack the range and complexity of human feeling demonstrated by Austen’s characters and understood by her readers.

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It used to be thought that the human brain could be neatly divided into these three distinct areas, a theory proposed by Paul MacLean. According to the theory of the triune brain, the brainstem forms the reptilian brain, the limbic areas form the mammalian brain, and the cortex forms the neo-mammalian brain. The brainstem is responsible for physiology; the mid-subcortical areas generate emotion; and the cortex orchestrates cognition, simple in most nonhuman animals and complicated in humans. MacLean’s triune brain makes sense in evolutionary terms since these areas did develop in this order. But his model is flawed in viewing these areas as functionally separate.

As subsequent research on the brain has demonstrated, neither emotions nor other kinds of thoughts and perceptions are processed in a discrete area of the brain. There’s no one emotional brain, as MacLean claimed, although we use the term as shorthand for all the systems that go into producing emotion. In the course of evolution, brain areas were not only added but integrated functionally in complex ways with structures that existed already. For instance, mammalian emotions involve input from the reptilian, physiological brain. There can’t be a separate emotion system, or any other kind of separate system, because all three “brains” are interconnected through neural circuitry that is astoundingly intricate, even in the lowliest mammals. I stress that the triune brain is an outmoded theory because it’s likely that every one of our responses, even simple ones, relies on parallel distributed processing, which means information flowing across the brain in all directions at any given moment. But triune brain theory nevertheless provides a useful way of thinking about the large divisions within brain anatomy.

The brain divides into halves as well as levels. There’s a right and a left hemisphere, familiarly known as the “left brain” and the “right brain.” These are for the most part symmetrical, with structures present in one half also present in the other. However, areas in each hemisphere differ to some extent in terms of function; this is known as laterality, and it’s seen throughout the animal kingdom although most pronounced in humans. In most humans, the capacity for language is located primarily in the cortex of the left hemisphere (you’ve already “met” Broca’s area), while awareness of the body and what it’s doing (called body maps) generally takes place in the cortex of the right hemisphere. A skilled writer and a graceful dancer—Jane was both—would need to have excellent capacities in both hemispheres. The “divide and conquer” structure of the brain’s anatomy enables different sides to specialize and so do a better job of their given tasks.

Asymmetry applies to subcortical areas as well; for instance, there’s a left- and a right-hemisphere amygdala. One of the functions of the amygdala is to decide what’s worth remembering, kind of like flagging a page of the novel you’re reading. One study concluded that, following the basic differences between the right and left hemispheres, the left amygdala might be more involved in helping us to remember emotional information that involves language whereas the right amygdala might help us to remember emotional information involving images. However, hemispheric distinctions are sometimes subtle or not entirely understood, and for this reason I’ll follow the lead of many books about the brain and refer to brain areas without differentiating between right and left hemispheres unless the distinction is relevant.

The cortex further subdivides into lobes, duplicated in each hemisphere; these are called the frontal, parietal, temporal, and occipital lobes. These lobes are carved out of the brain turned sideways. The frontal lobe, the foremost lobe of the brain, contains the motor cortex, which is responsible for initiating voluntary movement. The parietal lobe, on the top and middle of the brain, contains the somatosensory cortex, which governs the sense of touch. The temporal lobe, behind the ear, oversees hearing and language. The most forward area of the frontal lobe is appropriately named the prefrontal cortex. This is the “CEO” of the brain, responsible for executive functions such as planning and implementing behavior, problem solving, and logic. The visual cortex has its own lobe, the occipital lobe, which is located in the back of the brain. This is where Elizabeth’s image is assembled, in Darcy’s “mind’s eye.”

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TWO STAR PLAYERS: THE AMYGDALA AND THE ORBITOFRONTAL CORTEX (OFC)

This rough sketch of the brain doesn’t begin to capture its complexity, neither the extraordinary intricacy of its connections nor the abundance of areas that contribute to its functioning. But you don’t need to have a comprehensive knowledge of neuroanatomy to understand social intelligence, not even when that understanding incorporates a certain amount of neuroscience. In telling about some of the more physiological aspects of social intelligence, I’ve decided to let science take its cue from literature once again, using a version of the literary figure known as synecdoche.

Synecdoche is a rhetorical term like metaphor and simile. It most often means speaking of a part to refer to the whole, as in the phrase “hired hands” to mean workers. It’s the workers who are hired, not their hands, but hands stand in for the entire person. A synecdoche can also use the whole to represent a part, as in the phrase “the world isn’t treating you well.” Only a small portion of the world isn’t treating you well, and the rest is completely oblivious to your existence. In true synecdochal fashion, two brain areas will be referenced to discuss the bulk of information about the neurology of social intelligence: the amygdala and the orbitofrontal cortex, hereafter the OFC. (The OFC is also known as the orbital medial prefrontal cortex. According to some sources, the ventromedial prefrontal cortex [VMPFC] includes at least part of the OFC, but the boundaries of the VMPFC have not been firmly established.) The amygdala and the OFC are our “star players” when it comes to social intelligence.

This isn’t to say that other brain structures don’t play a vital role in social intelligence, or that they’ll be ignored, but these two areas will be featured because they are key to enabling our extraordinary sociality, greater than that of any other species. We say “hired hands” because the context in which the phrase developed concerned work done with the hands: carpentry, farming, manufacturing (the etymology of this last also refers to hands). We don’t say “hired feet” because the feet aren’t the important parts of the person who accomplishes such work. In the social brain, the amygdala and the OFC are significant hands, master builders of the social brain. If you become familiar with the OFC and the amygdala, you’ll not only know about two key areas, but you’ll also understand how regions of the brain connect and interact in a more general sense.

The OFC is a vital hub, where neural signals from far-flung areas are assembled. Although it’s technically part of the prefrontal cortex, the brain’s most advanced area, it’s located on the underbelly of the cortex, roughly behind your eyes. It sits at the juncture of limbic areas, which are crucially involved in the processing of emotion, and cortical areas, which enable cognition, including logic and reason. Among its many functions, it’s responsible for reading social signals, nonverbal expressions of emotion, intent, attitude, and so forth.

The brain is sometimes surprisingly literal in the sense that where something is located defines what it does (although, as you know, localized function acts in concert with information coming from many other brain areas). The location of the OFC has much to do with its function, for it coordinates information from cognitive and emotional areas. So crucial is the OFC’s role in emotional processing that although it’s technically considered part of the neocortex, it is more accurately defined as a cortico-limbic structure in terms of function as well as location. In addition, because it’s centrally located, more or less at the center of our brains, neural signals moving in every direction cross its path so that it receives information from every other system in the brain. It’s a key brain area for neural integration—the London of the brain.

What this means is that the OFC puts information from the external environment, some of which we’re conscious of, together with information from our internal environments, again some of which we recognize, such as conscious thoughts, but also endocrine, neurotransmitter, and hormone levels, which we know nothing about. Just as important as the information the OFC takes in is the information it sends out. The OFC has output connections to the amygdala and the hypothalamus, in charge of vital hormones, and to other brain areas where neurotransmitters are produced and released. It therefore orchestrates most of the important chemicals that control thought and behavior, the chemicals neurons need in order to know whether or not to be active. In short, the OFC is central in gathering information, interpreting perceptions, and generating responses.

The OFC is also connected to the brain’s reward system, also known as the seeking system. In the mind-brain sciences, reward refers to all the positive outcomes of actions and behavior, from applause after a presentation, to falling in love, to eating a chocolate bar. And speaking of chocolate, one study using fMRI (imaging of the brain in action) found that when people who habitually crave chocolate were presented with images of their favorite treat, their OFCs were more active than those of non-cravers. A related function, the OFC predicts and calculates the magnitude of reward and punishment, such as winning or losing money at gambling. People with damage to the OFC are poor gamblers.

Like all our brain and body parts, the OFC has a range of possible capability. The better the OFC functions, which depends primarily on the more connections it makes, the greater will be the flow of information throughout the brain. A high-functioning OFC leads to complexity. It gives us options and helps us make good choices. It does all this subconsciously, although its input contributes to conscious decision making. We can assume that Darcy’s OFC was higher functioning than that of Mr. Collins. Of course, we can say this about their brains as a whole, for Mr. Collins seems to be a very dim light indeed.

The amygdala is the gatekeeper for initiating stress reactions and other excitatory responses. It’s especially vital in the processing of fear and anxiety. It’s on the lookout for threats, so we can be prepared to confront them, and it activates when we encounter potentially dangerous situations or people, releasing chemical signals that get the body ready for action. It’s particularly receptive to visual information, which includes facial expression, so vital a sign for humans. I’m sure Darcy’s amygdala was highly active at the Meryton ball. But the amygdala is involved in positive responses as well, helping to generate the neurochemicals we need for excitement, which isn’t all that different from fear in terms of pure arousal. This explains why people can find fear and excitement difficult to separate, feeling a certain thrill, or “frisson,” as the French so elegantly say, in danger—think of riding a roller coaster.

As noted, the amygdala lets us know what’s worth paying attention to, and this again includes positive as well as negative experience. It tags people and events as important, and therefore worth remembering. Elizabeth remembers Darcy’s insult because of the amygdala, but this little structure also helps Jane to remember Bingley’s charming smiles. Like the OFC, the amygdala connects to areas such as the hypothalamus that control and produce vital neurochemicals. You can see an overlap in function with our two star players—the amygdala also interprets social signals, the OFC also notes what’s worth remembering, and both areas initiate neurochemical and physiological responses. But this isn’t unusual. Brain areas often duplicate one another in terms of function, varying somewhat in their activity to complement one another. Whatever our brains do, they usually require the coordination of many areas that contribute to or duplicate the same function.

The amygdala is under the jurisdiction of the OFC and must listen to its commands, which is reasonable since the OFC is receiving information from cortical-cognitive areas of the brain as well as subcortical-emotion areas. The amygdala, however, sends the OFC important information about the environment more quickly than cortical areas are capable of doing. Both structures, and indeed all brain areas, have input and output connections that involve them in continuous feedback loops that adjust our perceptions and responses millisecond by millisecond.

The OFC and amygdala pretty much run the social brain because working together, they initiate and regulate emotional responses. And emotions largely determine the nature of our relationships and interactions with others. You can be good at math without emotional input, but you need emotions to be good at relationships. This is ancient wisdom, common-sense “folk psychology.” But emotions are at the heart of other kinds of intelligence as well as social intelligence, which isn’t as obvious. Emotions are the prime movers of our mind-brains, as we see in the following chapter.

* We can assume that some of the land belonging to the estate is rented and that the family also has their own farm, which would provide them with milk, eggs, meat, fruits, and vegetables.

The notion of embodied cognition, that our bodies are central to thinking, has been advocated by other mind-brain scientists, although it doesn’t seem to budge the core belief that mind arises from brain. The logic of this conventional view is that even if events in the body affect the mind, such influence is still routed through the brain. Siegel disagrees about granting full primacy to this one organ.

Lewis Mehl-Madrona, Healing the Mind Through the Power of Story: The Promise of Narrative Psychiatry (Rochester, VT: Bear and Company, 2010), 13.

§ This is a reference to a thought experiment known as “the infinite monkey theorem,” the origins of which are unclear, but which was popularized in the early twentieth century by Émile Borel, a mathematician, and Sir Arthur Eddington, an astronomer. The premise is that monkeys, given an infinite amount of time in front of typewriters, would eventually type the works of Shakespeare, including Hamlet. Virtual monkeys sitting at virtual typewriters have nearly achieved this feat. In my view, using anything but real chimps and real typewriters (or, keeping up with the times, computers) is cheating!