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BIOSOCIAL APPROACHES In February of 1991, Stephen Mobley walked into a Domino’s Pizza store in Georgia to rob it. After getting the money, Mobley forced store manager John Collins onto his knees and shot him execution style. After committing several other robberies and bragging to friends about Collins’ murder, Mobley was apprehended by Atlanta police, charged with aggravated murder, and sentenced to death. In the automatic appeal to the Georgia Supreme Court to get his sentenced commuted to life in prison, his primary defense boiled down to claiming that his “genes made [him] do it.” In support of this defense, Mobley’s lawyers pointed to a Dutch study of an extended family in which, for generations, many of the men had histories of unpro- voked violence. The Researchers took DNA samples from 24 male members of the family and found that those with violent records had a marker for a mutant or vari- ation of a gene for the manufacture of monoamine oxydase (MAO), an enzyme that regulates a lot of brain chemicals. Mobley’s lawyers found a similar pattern of vio- lent behavior and criminal convictions among his male relatives across the genera- tions and asked the court for funds to conduct genetic tests on Mobley to see whether he had the same genetic variant. The court wisely denied the defense motion. Even if it were found that Mobley had the same genetic variant, it would not show that he lacked the substantial capacity to appreciate the wrongfulness of his acts or to conform to the require- ments of the law. Mobley’s lawyers were hoping to mitigate his sentence by appeal- ing to a sort of genetic determinism that simply does not exist. As we shall see in this section, genes don’t “make” us do anything; they simply bias us in one direc- tion rather than another. Except in cases of extreme mental disease or defect, we 273 S E C T I O N 8 08-Walsh (Introduction)-45498.qxd 2/13/2008 7:29 PM Page 273

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Page 1: BIOSOCIAL APPROACHES...BIOSOCIAL APPROACHES In February of 1991, Stephen Mobley walked into a Domino’s Pizza store in Georgia to rob it. After getting the money, Mobley forced store

BIOSOCIALAPPROACHES

In February of 1991, Stephen Mobley walked into a Domino’s Pizza store in Georgiato rob it. After getting the money, Mobley forced store manager John Collins ontohis knees and shot him execution style. After committing several other robberiesand bragging to friends about Collins’ murder, Mobley was apprehended by Atlantapolice, charged with aggravated murder, and sentenced to death. In the automaticappeal to the Georgia Supreme Court to get his sentenced commuted to life inprison, his primary defense boiled down to claiming that his “genes made [him] doit.” In support of this defense, Mobley’s lawyers pointed to a Dutch study of anextended family in which, for generations, many of the men had histories of unpro-voked violence. The Researchers took DNA samples from 24 male members of thefamily and found that those with violent records had a marker for a mutant or vari-ation of a gene for the manufacture of monoamine oxydase (MAO), an enzyme thatregulates a lot of brain chemicals. Mobley’s lawyers found a similar pattern of vio-lent behavior and criminal convictions among his male relatives across the genera-tions and asked the court for funds to conduct genetic tests on Mobley to seewhether he had the same genetic variant.

The court wisely denied the defense motion. Even if it were found that Mobleyhad the same genetic variant, it would not show that he lacked the substantialcapacity to appreciate the wrongfulness of his acts or to conform to the require-ments of the law. Mobley’s lawyers were hoping to mitigate his sentence by appeal-ing to a sort of genetic determinism that simply does not exist. As we shall see inthis section, genes don’t “make” us do anything; they simply bias us in one direc-tion rather than another. Except in cases of extreme mental disease or defect, we

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are always legally and morally responsible for our behavior. Cases such as Mobley’sunderline the urgent need for criminologists to understand the role of genes inhuman behavior as that role is understood by geneticists.

Biosocial criminologists believe that because humans have brains, genes, hormones,and an evolutionary history, they should integrate insights from the disciplines thatstudy these things into their theories and dismiss naïve nature versus nurture argu-

ments in favor of nature via nurture. Any trait, characteristic, or behavior of any living thingis always the result of biological factors interacting with environmental factors (Cartwright,2000), which is why we call modern biologically informed criminology biosocial rather thanbiological. Biosocial approaches have three broad complementary areas: behavior genetics,evolutionary psychology, and neuroscience.

y Behavior GeneticsBehavior genetics is a branch of genetics that studies the relative contributions of heredity andenvironment to behavioral and personality characteristics. Genes and environments work intandem to develop any trait—height, weight, IQ, impulsiveness, blood sugar levels, bloodpressure, and so on—the sum of which constitutes the person.

Behavior geneticists stress that genes do not cause us to behave or feel; they simply facili-tate tendencies or dispositions to respond to the environments in one way rather than inanother. There are no genes “for” criminal behavior, but there are genes that lead to particu-lar traits such as low empathy, low IQ, and impulsiveness that increase the probability of crim-inal behavior when combined with the right environments.

Behavior geneticists use twin and adoption studies to disentangle the relative influencesof genes and environments, and they tell us that genes and environments are always jointlyresponsible for any human characteristic. To ask whether genes or environment is mostimportant for a given trait is just as nonsensical as asking whether height or length is mostimportant to the area of a rectangle. Geneticists also tell us that gene expression depends onthe environment (think of identical rose seeds planted in an English garden and in the Nevadadesert, and then think about where the full potential of the seeds will be realized).

Behavior geneticists quantify the extent to which genes influence a trait with a measurecalled heritability (symbolized as h2), which ranges between 0 and 1. The closer h2 is to 1.0,the more the variance in a trait in a population, not in an individual, is due to genetic factors.Since any differences among individuals can only come from two sources—genes or environment—heritability is also a measure of environmental effects (1 – h2 = environmental effects). All cog-nitive, behavioral, and personality traits are heritable to some degree, with the traitsmentioned in the psychosocial section being in the .30 to .80 range (Carey, 2003).

Heritability estimates only tell us that genes are contributing to a trait; they do not tell uswhich genes; only molecular genetics can tell us this. Fortunately, we can now go beyond her-itability estimates and straight to the DNA by genotyping individuals using simple cheekswabs for about $10 each. This eliminates the need for special twin or adoptee samples, but wedo need cooperative studies with scientists having access to laboratories. Such studies arebeing conducted with increasing frequency, with the huge National Longitudinal Study of

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Adolescent Health (ADD Health) study being one inwhich some very important genetic findings (the dis-covery of specific genes) relating to criminal behaviorhave been recorded (Beaver, 2006). It is emphasized thatany individual gene only accounts for a miniscule pro-portion of the variance in criminal behavior, and that itcontributes to a trait linked to criminality, not to crim-inality itself. Genes always have indirect effects onbehavior via their effects on traits.

y Gene/Environment Interaction and Correlation

Gene/environment (G/E) interaction and G/E correla-tion describe people’s active transactions with theirenvironment. G/E interaction involves the notion thatpeople are differentially sensitive to identical environ-mental influences and will thus respond in differentways to them. For instance, a relatively fearless andimpulsive child is more likely to seize opportunities toengage in antisocial behavior than is a fearful and con-strained child. G/E correlation means that genotypesand the environments they find themselves in arerelated. These concepts enable us to conceptualize theindirect way that genes help to determine what aspectsof the environment will and will not be important to us.There are three types of G/E correlation: passive, reac-tive, and active.

Passive G/E correlation is the positive associationbetween genes and their environments that existsbecause biological parents provide children with genesfor certain traits and an environment favorable for theirexpression. Children born to intellectually giftedparents, for instance, are likely to receive genes forabove-average intelligence and an environment inwhich intellectual behavior is modeled and reinforced,thus setting them on a trajectory independent (pas-sively) of their actions.

Reactive G/E correlation refers to the way othersreact to the individual on the basis of his or her evoca-tive behavior. Children bring traits with them to situa-tions that increase or decrease the probability of evoking certain kinds of responses fromothers. A pleasant and well-mannered child will evoke different reactions than will a bad-tempered and ill-mannered child. Some children may be so resistant to socialization thatparents may resort to coercive parenting or simply give up, either of which may worsen any

▲ Photo 8.1 Former major league baseballplayer Jose Canseco is sworn in at a U.S. House ofRepresentatives baseball steroids hearing. Cansecopresents a fascinating case for biosocial theories.Jose had a fraternal twin brother, Ozzie, who alsochose a career in baseball. However, in comparisonwith Jose’s 462 home runs and over 1,400 RBI,Ozzie had only a “cup of coffee” in the majorleagues. He came to bat only 65 times over 3seasons and never hit a home run. Had he been anidentical rather than a fraternal twin, might Ozziehave performed more like his brother? Afterfinishing his baseball career, Jose wrote a book(Juiced) in which he admitted using steroids formost of his playing career and claimed that 85% ofother players in his era did likewise. Because of hissteroid use, many baseball experts predict Jose willnever be elected to the baseball Hall of Fame,though his career numbers exceed those of manycurrent Hall of Fame players.

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antisocial tendencies and drive them to seek environments where their behavior is accepted.Reactive G/E correlation thus serves to magnify phenotypic differences by funneling individ-uals into like-minded peer groups (“birds of a feather flock together”).

Active G/E correlation refers to the active seeking of environments compatible with ourgenetic dispositions. Active G/E correlation becomes more pertinent as we mature and acquirethe ability to take greater control of our lives because within the range of possibilities avail-able in our cultures, our genes help to determine what features of the environment will andwill not be attractive to us. Our minds and personalities are not simply products of externalforces, and our choices are not just passive responses to social forces and situations. We areactive “niche picking” agents who create our own environments just as they help to create us.

Genes imply human self-determination because, after all, our genes are our genes. AsColin Badcock (2000) put it: “Genes don’t deny human freedom; they positively guarantee it”(p. 71). Genes are constantly at our beck and call, extracting information from the environ-ment and manufacturing the substances we need to navigate it. They are also what make usuniquely ourselves and thus resistant to environmental influences that grate against ournatures. In short, genes do not constrain us, they enable us. This view of humanity is far morerespectful of human dignity than the blank slate view that we are putty in the hands of theprevailing environmental winds.

y Behavior Genetics and Criminal BehaviorA review of 72 behavior genetic studies conducted up to 1997 found that 93% were support-ive of the hypothesis than genes affect antisocial behavior (Ellis & Walsh, 2000). Althoughthere are no behavior genetic theories of criminal behavior per se, behavior genetic studieshelp us to temper and make more sense of traditional criminological theories. For instance,large behavior genetic studies conducted in the U. S. (Cleveland, Wiebe, van den Oord, &Rowe, 2000) and the U. K. (Moffitt & The E-Risk study team, 2002) showed that genetic fac-tors play a large part in sorting individuals into different family structures (broken vs. intacthomes), a variable often linked to antisocial behavior.

A major longitudinal study of child abuse and neglect that integrated genetic (the sameMAO gene mentioned in the section vignette) data showed why only about one-half ofabused/neglected children become violent adults (Caspi et al., 2002). This study showed thatneither genetic risk nor abuse/neglect by themselves have much effect on antisocial behavior,but the odds of having a verified arrest history for a violent crime for subjects with bothgenetic and environmental risk factors (G/E interaction) were 9.8 times greater than the oddsfor subjects with neither.

Another study looked at Gottfredson and Hirschi’s (1990) assumption that parents areprimarily responsible for their children’s self-control (Wright & Beaver, 2005). A modest rela-tionship between parental practices and children’s self-control was found, but disappearedwhen genetic information was added. In other words, not using genetically informed methodsleads researchers to misidentify important causal influences. Wright and Beaver (2005) con-cluded, “for self-control to be a valid theory of crime it must incorporate a more sophisticatedunderstanding of the origins of self-control” (p. 1190).

Unlike the relatively strong genetic influences discovered for most human traits, geneticinfluences on antisocial behavior are modest, especially during the teenage years. A study of3,226 twin pairs that found genes accounted for only 7% of the variance in antisocial behavior

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among juvenile offenders, but 43% among adult offenders (Lyons et al., 1995). Heritabilitycoefficients for most traits related to antisocial behavior are typically in the .20 to .80 range,and for antisocial behavior itself, two meta-analyses concluded that they are in the .40 to .58range (Miles & Carey, 1997; Rhee & Waldman, 2002), with h² being higher in adult than injuvenile populations.

What this means is that the majority of delinquents have little, if any, genetic vulnerabil-ity to criminal behavior while a small minority may have considerable vulnerability. Poolingthese two groups has the effect of elevating estimates of the overall influence of genes whileminimizing it for those most seriously involved. For instance, although Mednick, Gabrielli,and Hutchings (1984) found a weak overall pattern of genetic effects for delinquency amonga large number of young males, just 1.0% of the cohort (37 males) who had biological fatherswith three or more criminal convictions accounted for fully 30% of all the cohort’s convic-tions. Genetic effects on antisocial behavior are most likely to be found among chronic offend-ers who begin offending prior to puberty and who continue to do so across the life course(Moffitt & Walsh, 2003).

The article by Anthony Walsh in this section further explains behavior genetic conceptsand ties them to the anomie/strain theory. Walsh contends that achieving the AmericanDream, the success goal so central to anomie/strain theory, is a matter of having the requisitecognitive and temperamental characteristics to perform well in school and at work. He showshow behavior genetics can augment sociological criminology theories by illuminating thegenetic underpinnings of many of their favored concepts such as SES, occupational mobility,frustration, and so forth.

Evolutionary PsychologyEvolutionary psychology explores human behavior using an evolutionary theoretical frame-work and seeks to explain human behavior with reference to human evolutionary history.Criminologists operating within the evolutionary framework explore how certain behaviorssociety now calls criminal may have been adaptive in ancestral environments. Evolutionarypsychology complements behavior genetics because it informs us how the genes of interestcame to be present in the first place. The primary difference between the two disciplines is thatwhile behavior genetics looks for what makes people different, evolutionary psychology focuseson what makes us all the same. Another basic difference is that evolutionary psychology looksat ultimate level “why” questions (what evolutionary problem did this behavioral mechanismevolve to solve?), and behavioral geneticists look at proximate level “how” questions (to whatextent is this behavioral mechanism influenced by genes in this population at this time?).

Evolutionary psychologists agree with most criminologists that although it is morallyregrettable, crime is normal behavior for which we all have the potential (Kanazawa, 2003).Evolutionary logic tells us that if criminal behavior is normal, it must have conferred some evo-lutionary advantage on our distant ancestors. However, because modern environments are soradically different from the hunter/gatherer environments in which we evolved, many traitsselected for their adaptive value at the time may not be adaptive today. It is important to real-ize that it is the traits underlying criminal behavior that are the alleged adaptations, not the spe-cific acts; genes do not code themselves for burglarizing a house or stealing a car (Rowe, 1996).

Criminal behavior is a way to acquire resources illegitimately. Evolutionary psycholo-gists refer to such behavior (whether it is defined as criminal or not) as cheating, and think

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of individual traits associated with it, such as impulsiveness, aggression, and low empathy,in terms of normal distributions dispersed around adaptive species averages. Whetherexploitation occurs depends on environmental triggers interacting with individual differ-ences and with environmental constraints. Although we all have the potential to exploit anddeceive others, we are a highly social and cooperative species with minds forged by evolu-tion to form cooperative relationships built on trust (Barkow, 2006). Cooperation is typi-cally contingent on the reciprocal cooperation of others, and is thus a tit-for-tat strategyfavored by natural selection because of the benefits it confers. We cooperate with our fellowsbecause we feel good when we do and because it identifies us as reliable and trustworthy,which confers valued social status on us.

Because cooperation occurs among groups of other cooperators, it creates niches for non-cooperators to exploit others by signaling their cooperation and then failing to follow through(Tibbetts, 2003). In the human species, criminal behavior may be viewed as an extreme formof defaulting on the rules of cooperation. But cheating comes at a cost, so before deciding todo so the individual must weigh the costs and benefits of cooperating versus defaulting.Cheating is rational (not to be confused with moral) when the benefits outweigh the costs. Butif cheating is so rational, how did cooperation come to be predominant in social species? Theanswer is that cheating is only rational in circumstances of limited interaction and communi-cation. Frequent interaction and communication breeds trust and bonding, and cheatingbecomes a less rational strategy because cooperators remember and retaliate against thosewho have cheated them. Ultimately, cooperation is the most rational strategy in any socialspecies because each player reaps in the future what he or she has sown in the past.

Yet, we continue to see cheating behavior despite threats of exposure and retaliation. Wedo so because exposure and retaliation are threats only if cheats are constrained to operatewithin the same environment in which their reputations are known. Cheats can move fromlocation to location meeting and cheating a series of others who are unaware of their reputa-tion. This is the pattern of many career criminals who move from place to place, job to job,and relationship to relationship, leaving a trail of misery behind them before their reputationcatches up. This is why cheats are more likely to prosper in large cities in modern societies thanin small traditional communities where the threat of exposure and retaliation is great (Ellis &Walsh, 1997).

y The Evolution of Criminal TraitsThere are a number of evolutionary theories of crime, all of which focus on reproductivestrategies. This is not surprising because from a biological point of view, the evolutionaryimperative of all living things is reproductive success. There are two ways that members of anyanimal species can maximize reproductive success: parenting effort and mating effort.Parenting effort is the proportion of reproductive effort invested in rearing offspring, andmating effort is that proportion allotted to acquiring sexual partners. Because humans areborn more dependent than any other animal, parenting effort is particularly important to ourspecies. The most useful traits underlying parenting effort are altruism, empathy, nurturance,and intelligence (Rowe, 2002).

Humans invest more in parenting effort than any other species, but there is considerablevariation within the species. Gender constitutes the largest division due to different levels ofobligatory parental investment between the genders. Female parental investment necessarily

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requires an enormous expenditure of time and energy, but the only obligatory investment ofmales is the time and energy spent copulating. Reproductive success for males increases inproportion to the number of females to whom they have sexual access, and thus males haveevolved a propensity to seek multiple partners. Mating effort emphasizes quantity over qual-ity (maximizing the number of offspring rather than nurturing a few), although maximizingoffspring numbers is obviously not a conscious motive of any male seeking sex. The proximatemotivation is sexual pleasure, with more offspring being a natural consequence (in pre-contraceptive days) when the strategy proved successful.

Reproductive success among our ancestral females rested primarily on their ability tosecure mates to assist them in raising offspring in exchange for exclusive sexual access, andthus human females evolved a much more discriminating attitude about sexual behavior(Fisher, 1998; Geary, 2000). According to evolutionary biologists, the inherent conflict betweenthe reckless and indiscriminate male mating strategy and the careful and discriminatingfemale mating strategy drove the evolution of traits such as aggressiveness and the lowering oftrait levels (relative to female levels) such as empathy and constraint that help males to over-come both male competitors and female reticence. The important point to remember is thatalthough these traits were designed by natural selection to facilitate mating effort, they are alsouseful in gaining non-sexual resources via illegitimate means (Quinsey, 2002; Walsh, 2006).

The reverse is also true—traits that facilitate parenting effort underlie other forms ofprosocial activity: “Crime can be identified with the behaviors that tend to promote matingeffort and noncrime with those that tend to promote parenting effort” (Rowe, 1996, p. 270).Because female reproductive success hinges more on parenting effort than mating effort,females have evolved higher levels of the traits that facilitate it (e.g., empathy and altruism)and lower levels of traits unfavorable to it (e.g., aggressiveness) than males. Of course, bothmales and females engage in both mating and parenting strategies, and both genders follow amixed mating strategy. It is only claimed that mating behavior is more typical of males andparenting effort is more typical of females.

Empirical research supports the notion that an excessive concentration on mating effortis linked to criminal behavior. A review of 51 studies relating number of sex partners to crim-inal behavior found 50 of them to be positive, and in another review of 31 studies it was foundthat age of onset of sexual behavior was negatively related to criminal behavior (the earlier theage of onset, the greater the criminal activity) in all 31 (Ellis & Walsh, 2000). A British cohortstudy found that the most antisocial 10% of males in the cohort fathered 27% of the children(Jaffee, Moffitt, Caspi, & Taylor, 2003), and anthropologists tell us that there are striking dif-ferences in behavior between members of cultures that emphasize either parenting or matingstrategies. The world over, cultures emphasizing mating effort exhibit behaviors (low-levelparental care, hypermasculinity, transient bonding) considered antisocial in Western societies(Ember & Ember, 1998). Finally, a recent molecular genetic study found the same genes thatwere significantly related to number of sexual partners were also significantly related to anti-social behavior (Beaver, Wright, & Walsh, in press).

y The NeurosciencesWhatever the source of human behavior, it is necessarily funneled through the brain,arguably the most awe-inspiring structure in the universe. Although the brain is only about2% of body mass, it consumes 20% of the body’s energy as it perceives, evaluates, and

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responds to its environment (Shore, 1997). This three-pound marvel of evolutionarydesign is the CEO of all that we think, feel, and do. All our thoughts, feelings, emotions,and behavior are the results of communication networks of brain cells called neurons. Themore “primitive” networks come “hard wired” at birth, but development of the higherbrain areas depends a lot on environmental “software” downloaded after birth. The mes-sage neuroscience has for us is that the experiences we encounter largely determine the pat-terns of our neuronal connections, and thus our ability to successfully navigate our lives(Quartz & Sejnowski, 1997).

Neural networks are continually being made and selected for retention or elimination ina use-dependent process governed by the strength and frequency of experience, and it is biasedin favor of networks that are most stimulated during early development (Restak, 2001). Thisis why bonding and attachment are so vital to human beings, and why abuse and neglect is soinjurious. Chronic stress can produce neuron death via the production of stress hormones,and children with high levels of these hormones experience cognitive, motor, and social devel-

opment delays (Gunnar, 1996). As Perry andPollard (1998) point out: “Experience in adultsalters the organized brain, but in infants andchildren it organizes the developing brain” (p. 36).Brains organized by stressful and traumatic eventstend to relay events along the same neural path-ways laid out by those early events because path-ways laid down early in life are more resistant toelimination than pathways laid down later in life.A brain organized by negative events is ripe forantisocial behavior.

280 INTRODUCTION TO CRIMINOLOGY

▲ Photo 8.2 Harkening back to the 19th century, whenpostmortem examinations of the brains of criminals were afrequent phenomenon, the brain of serial killer John WayneGacy was dissected after his execution. The attempt to locatean organic explanation of his monstrous behavior wasunsuccessful.

y Reward Dominance andPrefrontal DysfunctionTheories

Reward dominance theory is a neurologicaltheory based on the proposition that behavior is regulated by two opposing mechanisms, thebehavioral activating system (BAS) and thebehavioral inhibition system (BIS). The BAS isassociated with the neurotransmitter dopamineand with pleasure areas in the brain (Gove &Wilmoth, 2003). The BIS is associated with sero-tonin and with brain structures that govern mem-ory (Pinel, 2000). Neurotransmitters such asdopamine and serotonin are the chemical mes-sengers that shunt information between neuralnetworks. Dopamine facilitates goal-directedbehavior and serotonin generally modulatesbehavior (Depue & Collins, 1999).

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The BAS is sensitive to reward and can be likened to an accelerator motivating a personto seek rewarding stimuli. The BIS is sensitive to threats of punishment and can be likened toa brake that stops a person from going too far too fast. The BAS motivates us to seek whateveraffords us pleasure, and the BIS tells us when we have had enough for our own good. A nor-mal BAS combined with a faulty BIS, or vice versa, may lead to a very impulsive person witha “craving brain” that can lead him or her into all sorts of physical, social, moral, and legal dif-ficulties, by becoming addicted to pleasures such as food, gambling, sex, alcohol, and drugs(Ruden, 1997).

While most of us are more or less equally sensitive to both reward and punishment (instate of dopamine/serotonin balance), in some people one system might dominate the othermost of the time (Ruden, 1997). The theory asserts that criminals, especially chronic crimi-nals, have a dominant BAS, which tends to make them overly sensitive to reward cues and rel-atively insensitive to punishment cues (Lykken, 1995). Reward dominance theory provides uswith hard physical evidence relating to the concepts of sensation seeking, impulsiveness, andlow self-control we have previously discussed because each of these traits is underlain by eithera sticky accelerator (high dopamine) or faulty brakes (low serotonin).

A third system of behavior control is the flight/fight system (FFS) chemically controlledby epinephrine (adrenaline). The FFS is that part of the autonomic nervous system that mobi-lizes the body for vigorous action in response to threats by pumping out epinephrine. Fear andanxiety at the chemical level is epinephrine shouting its warning: “Attention, danger ahead;take action to avoid!” Having a weak FFS (low epinephrine) that whispers rather than shouts,combined with a BAS (high dopamine) that keeps shouting “Go get it” and a BIS (low sero-tonin) too feeble to object, is obviously very useful when pursuing all kinds of criminal andantisocial activities.

Another neurologically specific theory of criminal behavior is prefrontal dysfunctiontheory. The prefrontal cortex (PFC) is responsible for a number of human attributes such asmaking moral judgments, planning, analyzing, synthesizing, and modulating emotions. ThePFC provides us with knowledge about how other people see and think about us, thus mov-ing us to adjust our behavior to consider their needs, concerns, and expectations of us. ThesePFC functions are collectively referred to as executive functions and are clearly involved inprosocial behavior. If these functions are compromised in some way via damage to the PFC,the result is often antisocial behavior.

Positron emission tomography (PET) and functional magnetic resonance imaging(fMRI) studies consistently find links between PFC activity and impulsive criminal behavior.A PET study comparing impulsive murderers with murderers whose crimes were plannedfound that the former showed significantly lower PFC and higher limbic system activity(indicative of emotional arousal) than the latter and other control subjects (Raine, Meloy,Bihrle, Stoddard, LaCasse, & Buchsbaum, 1998). Cauffman, Steinberg, and Piquero (2005)combined reward dominance and PFC dysfunctions theories in a large-scale study of incar-cerated and nonincarcerated youths in California and found that seriously delinquent offend-ers have slower resting heart rates and performed poorly relative to nondelinquents on variouscognitive functions mediated by the PFC.

Jana Bufkin and Vickie Lettrell’s review of 17 neuroimaging studies in this section showconclusively that impulsive violent behaviors are associated with PFC deficits. They interpretthe findings in terms of problems with the PFC’s regulation of negative emotionality. Theyalso explore activity between the PFC and subcortical structures associated with violence and

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aggression. Like many biosocial researchers, they make a plea for interdisciplinary studies ofcriminality so that the biological, psychological, and social factors associated with it can bebetter understood.

Lee Ellis’s evolutionary neuroandrogenic (ENA) theory presented in this sectionattempts to tie many genetic, evolutionary, and neurological factors together into a unifiedtheory. Ellis claims that his theory can explain why many of the demographic correlates ofcrime, such as age, sex, and SES, as well as biological correlates, such as mesomorphic bodybuild, heart rate, and brainwave patterns, exist. He begins with two assumptions that areultimately about mating versus parenting effort: (1) Males have been naturally selected forengaging in status striving and resource procurement because over countless generations,females who have chosen mates based on a male’s ability to obtain resources will have leftmore offspring in subsequent generations than females who use other criteria for selectingmates. (2) Fetal exposure of male brains to testosterone makes them more prone to compet-itive status striving than females.

Table 8.1 summarizes the strengths and weaknesses of biosocial perspectives andtheories.

y Evaluation of the Biosocial PerspectiveBiosocial theories have never been popular with mainstream social scientists because theywere interpreted as implying a Lombrosian biological inferiority of criminals. This kind ofthinking is rarer today as social scientists have become more sophisticated about the inter-action of biology and environment (Robinson, 2004). There are still people who fear that“biological” theories can be used for racist ends, but as Bryan Vila (1994) remarks, “Findingscan be used for racist or eugenic ends only if we allow perpetuation of the ignorance thatunderlies these arguments” (p. 329). Bigots and hate-mongers will climb aboard any vehiclethat gives their prejudices a free ride, and they have done so for centuries before genes wereheard of.

The strength of biosocial approaches lies in their ability to incorporate biological factorsinto their theories and to physically measure many of them via various chemical, electrophys-iological, and neuroimaging methods. However, studies are more difficult to do and far moreexpensive than the typical social science study. If we want genetic information, we cannotsimply go to the nearest high school and survey a few hundred students. Genetic studiesrequire comparing samples consisting of pairs of identical and fraternal twins and/oradoptees, and these are difficult to come by. However, new technologies have allowed us to gostraight to the DNA, thus eliminating the need for special samples consisting of pairedsubjects with known degrees of genetic relatedness.

It is difficult to make generalizations from the typical neuroimaging study because manyconsist of a small number of known offenders matched with a control group. Neuroimagingstudies are extremely expensive to conduct (an average of $3,000 per scan), and many of themmay only consist of 20 or 30 identified criminals matched with a control group of similar size.Nevertheless, biosocial studies provide criminologists with “harder” evidence than they aretypically able to get, and this evidence will help them to more solidly ground their theories,and when numerous small studies tell the same story we are able to place more confidence intheir conclusions.

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Section 8 � Biosocial Approaches 283

Table 8.1 Summarizing Biosocial Perspectives and Theories

Theory

BehaviorGeneticsPerspective

EvolutionaryPsychologyPerspective

NeurosciencePerspective

RewardDominanceTheory

PrefrontalDysfunctionTheory

Key Concepts

Genes affect behavior ininteraction with environmentalinfluences. Heritability estimatesthe relative contribution ofgenetic and environmentalfactor traits affecting criminality.All individual traits are at leastmodestly influenced by genes.

Human behavior is rooted inevolutionary history. Naturalselection has favored victimizingtendencies in humans, especiallymales. These tendencies arose tofacilitate mating effort, but areuseful in pursuing criminalbehavior as well. Criminalsemphasize mating effort overparenting effort more thanmales in general.

Whatever its origin, all stimuliare channeled through the brainbefore being given expression inbehavior. The development of thebrain is strongly influenced byearly environmental experiences,especially those involvingnurturance and attachment.

Behavioral activating (BAS) andbehavioral inhibiting system(BIS) are dopamine andserotonin driven, respectively.Among criminals the BAS tendsto be dominant over the BIS.This BIS/BAS imbalance can leadto addiction to many things,including crime.

Frontal lobes control long-termplanning and temper emotionsand their expressions. Criminalshave frontal lobes that failto function as they do inmost people, especially interms of inhibiting actionsthat harm others.

Strengths

Looks at both the genetic andenvironmental risk factors forcriminal behavior. Understandinggenetic contributions alsoidentifies the complementarycontributions of environmentalfactors.

Ties criminology to evolutionarybiology. Mating effort helps toexplain why males are morecriminal than females and whycriminals tend to be moresexually promiscuous thanpersons in general. Emphasizesthat crime is biologically“normal” (although regrettable)rather than pathological.

Shows how environmentalexperiences are physically“captured” by the brain.Emphasizes the importance ofnurturing for optimaldevelopment of the brain. Usessophisticated technology andprovides “harder” evidence.

Explains why low serotoninis related to offending (lowserotonin = low self-control).Explains why criminality ispersistent in some offendersbecause they develop a taste for the “thrill of it all.”

Explains why moral reasoning isinversely related to involvementin persistent criminalityExplains why criminality hasbeen linked to frontal lobedamage and to abnormalbrain waves.

Weaknesses

Requires samples of twinsand/or adoptees, which aredifficult to come by. However,technology now enables us toeliminate the need for specialsamples and go straight to theDNA.

Gives some the impression thatbecause crime is considered“normal” it is justified orexcused. Makes assumptionsabout human nature which mayor may not be true. Whilerecognizing that cultureis important, it tends to ignore it.

High cost of neuroimagingstudies. Very small samplesof known criminals are used,thus limiting generalizations.Linking specific brain areas tospecific behaviors isproblematic.

The neurological underpinningsof the BAS and BIS have beendifficult to precisely identify.Studies difficult and expensiveto conduct.

Dysfunction of the prefrontallobes remains difficult tomeasure, even with fMRI scans.Some sampling difficultiesnoted for the neurosciences ingeneral.

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y Policy and Prevention:Implications of Biosocial Theories

The policy issues suggested by the biosocial perspective are midway between the macro-levelsociological suggestions aimed at whole societies or communities and the micro-level sugges-tions of psychological theories aimed at already convicted criminals. Mindful of how nurtur-ing affects both gene expression and brain development in humans, many biosocialcriminologists have advocated a wide array of “nurturant” strategies such as pre- and postna-tal care for all women, monitoring infants and young children through the early developmen-tal years, paid maternal leave, nutritional programs, and a whole host of other interventions(Vila, 1997; Walsh & Ellis, 1997).

Biosocial criminologists are typically in the forefront in advocating treatment over punish-ment, and toward this end, they have favored indeterminate sentences over fixed sentences(Lanier & Henry, 1998). Pharmacological treatments in conjunction with psychosocial treat-ments have proven to be superior to psychosocial treatment alone for syndromes (alcoholism,drug addiction, etc.) associated with criminal behavior (Walsh, 2002). Of course, there arealways dangers of seeking simple medical solutions to complex social problems. Requiring sexoffenders to take anti-androgen treatment to reduce the sex drive raises both medical andlegal/ethical issues regardless of how effective the treatment is. Prescribing selective serotoninreuptake inhibitors such as Prozac and Zoloft helps to curb low self-control and irritability, butthere is always the temptation to treat everyone the same regardless of their serotonin levels.

Some behavioral scientists tend to feel that identifying biological risk factors will lead tocessation of efforts to reduce crime through environmental improvement because such factorsare wrongly thought to be intractable. But biosocial studies provide information about bothenvironmental and biological risk factors and, as such, are “more likely to refine social policiesby better specification of environmental factors than to divert funds from environmentalcrime prevention strategies” (Morley & Hall, 2003). In other words, they will enable us tobetter pinpoint environmental factors that may prove fruitful in our crime prevention efforts.

y Summary◆ Behavior geneticists study the genetic underpinning of traits and characteristics in

populations by calculating heritability coefficients. There are no genes “for” any kind ofcomplex human behavior; genes simply bias trait values in one direction or another. Thisview is respectful of human dignity because it implies self-determinism because our genesare our genes.

◆ Gene/environment interaction tells us that the impact an environmental situation(e.g., living in a crime ridden neighborhood) has on us depends on who we are, andgene/environment correlation tells us that who we are is a product of our unique genotypeand the environments we find ourselves in.

◆ Genes have practically no influence on juvenile delinquency, probably because of thehigh base rate of delinquency. There are genetic effects for chronic and serious delinquents,but these few individuals tend to get “lost” in studies that combine them with those who limittheir offending to adolescence. Adult criminality is much more influenced by genes. One of

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Section 8 � Biosocial Approaches 285

the reasons that we find only modest genetic effects in criminality when the traits that under-lie it are strongly influenced by genes is that parents have control over their children’s behav-ior, but little or none over the underlying traits.

◆ Evolutionary psychology focuses on why we have the traits we do and is more inter-ested in their universality than in their variability. Crime is viewed as a normal but regrettableresponse to environmental conditions. By this it is meant that many human adaptationsforged by natural selection in response to survival and reproductive pressures are easily co-opted to serve morally wrong purposes.

◆ In common with all sexually producing species, humans are preeminently concernedwith their own survival and reproductive success. The traits designed to assist males in theirmating efforts include many that can also assist them to secure other resources illegitimately;traits designed to assist females in their parenting efforts are conducive to prosocial behavior.Mating vs. parenting effort is not an either/or thing; males and females engage in both at var-ious times in their lives, it is just that mating effort is more typical of males and parentingeffort is more typical of females

◆ Socially cooperating species create niches that cheats can exploit to their advantage bysignaling cooperation but then defaulting. Cheating is a rational strategy in the short term, butinvites retaliation in the long term. This is why chronic criminals rarely have successful rela-tionships with others and why they typically die broke.

◆ Neuroscience tells us that genes have surrendered control of human behavior to thebrain. Following genetic wiring to jump-start the process, the brain literally wires itself inresponse to environmental input. The softwiring of our brains is an electro-chemical processthat depends on the frequency and intensity of early experiences. Adverse experiences can lit-erally physically organize the brain so that we experience the world negatively, which is whynurturing, love, and attachment are so important to the healthy development of humans.

◆ Reward dominance theory informs us that the brain regulates our behavior throughthe BIS and BAS systems. The BIS and BAS systems (underlain by serotonin and dopamineneurotransmitters, respectively) in most people are balanced, but criminals tend to have eitheran overactive BAS or an underactive BIS. This means that their behavior is dominated byreward cues and relatively unaffected by punishment cues.

◆ Prefrontal dysfunction theory posits that the brain’s prefrontal cortex (PFC) is vital tothe so-called executive functions such as planning and modulating emotions. If the PFC isdamaged in any way, the individual is deficient in these executive functions and tends to beimpulsive.

E X E R C I S E S A N D D I S C U S S I O N Q U E S T I O N S

1. If it could be shown with high scientific confidence that some young children inheritgenes that put them at 85% risk for developing antisocial proclivities, what do youthink should be done? Should their parents be warned to be especially vigilant and toseek early treatment for their children, or would such a warning tend to stigmatizechildren? What are the costs and benefits of each option?

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2. We know that males, especially young males, are more likely to perpetrate and be vic-timized by violent crimes. Provide a plausible evolutionary explanation for this.

3. How might reward dominance theory add strength and coherence to low self-controltheory?

U S E F U L W E B S I T E S

Behavioral Genetics. http://www.ornl.gov/sci/techresources/Human_Genome/elsi/behavior.shtml.

Crime Times. http://crimetimes.org/.

Evolutions Voyage. (Evolutionary Psychology). http://www.evoyage.com/.

The Human Brain. http://www.fi.edu/brain/index.htm.

Naturalistic Fallacy. http://www.iscid.org/encyclopedia/Naturalistic_Fallacy.

Anatomy of the Brain. http://www.neuroguide.com/index.html.

G L O S S A RY

Behavior genetics: A branch of genetics that studies the relative contributions of heredity andenvironment to behavioral and personality characteristics.

Behavioral activating system: A brain reward system associated chemically with the neuro-transmitter dopamine.

Behavioral inhibition system: A brain system that inhibits or modulates behavior and is associated with serotonin.

Evolutionary psychology: A way of thinking about human behavior using a Darwinian evolutionary theoretical framework.

Flight/fight system: An autonomic nervous system mechanism that mobilizes the body foraction in response to threats by pumping out epinephrine.

G/E correlation: The notion that genotypes and the environments they find themselves in arerelated because parents provide children with both.

G/E interaction: The interaction of genes and the environment.

Heritability: A concept defined by a number ranging between 0 and 1 indicating the extent towhich variance in a phenotypic trait in a population is due to genetic factors.

Mating effort: The proportion of total reproductive effort allotted to acquiring sexual part-ners; traits facilitating mating effort are associated with antisocial behavior.

Neurons: Brain cells consisting of the cell body, an axon, and a number of dendrites.

Neurotransmitters: Brain chemicals that carry messages from neuron to neuron.

Parenting effort: The proportion of total reproductive effort invested in rearing offspring;traits facilitating parenting effort are associated with prosocial behavior.

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Section 8 � Behavior Genetics and Anomie/Strain Theory 287

Prefrontal cortex: Part of the brain that plays the major integrative and supervisory roles inthe brain.

Prefrontal dysfunction theory: A neurological theory of antisocial behavior based on dys-function of the prefrontal cortex.

Reward dominance theory: A neurological theory based on the proposition that behavior isregulated by two opposing mechanisms, the behavioral activating system (BAS) and thebehavioral inhibition system (BIS).

SOURCE: Walsh, A. (2000). Behavior genetics and anomie strain theory. Criminology, 38(4). Reprinted with permission of theAmerican Society of Criminology.

READING

Behavior Genetics and Anomie/Strain Theory

Anthony Walsh

In this article, Anthony Walsh states that criminology is in need of a conceptual boost from themore fundamental sciences such as behavior genetics. He claims that behavior genetics is a bio-logically friendly environmental discipline because it tells us just as much about the effect of theenvironment on a trait as it does that of genes. He uses anomie/strain theory to illustrate this.Anomie/strain is about occupational success, or the lack thereof, and Walsh looks at the role ofintelligence and temperament, both of which are highly heritable, on occupational success. Healso shows how behavior genetics can throw light on many of criminology’s favored concepts.

The decade of the 1990s saw numerous reports ofcrisis within sociology in general as well as incriminology in particular. The substance of mostof these reports is that the discipline is moribundand desperately needs something to breathe freshlife into it. The reason most often given for thisstate of affairs is sociology’s dogged refusal toentertain the possibility that biological factorsmay play an important role in helping us tounderstand phenomena of concern to us.Perhaps a major reason for this refusal can befound in what is probably the most cited passagein all of sociology: “The determining cause of asocial fact should be sought among antecedentsocial facts and not among the states of the

individual consciousness” (Durkheim, 1982:134).This dictum has become a sociological mantraused to assert and defend the ontological autonomyof the discipline. According to Udry (1995:1267),however, although Durkheim clearly meant thisstatement to be a boundary axiom defining soci-ology’s purview, sociologists came to think of itas “a true statement about the nature of the worldinstead of a set of deliberate blinders to helpthem focus their attention.”

There are signs that certain criminologicaltheories are becoming more psychologically andbiologically informed, and there have been anumber of recent encapsulations of biosocialapproaches in the criminology literature. Authors

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of such overviews, however, tend not to be main-stream criminologists, but psychologists, psychi-atrists, and biologists, and none of them has triedto show the relevance of biosocial variables tomainstream criminological theories. Perhapsbiologically informed theories will continue tohave minimal impact on mainstream criminol-ogy until it can be shown that they are not anti-thetical to current environmental theories ofcrime. My primary purpose in this paper is to demonstrate that at least one biologicallyinformed discipline––behavior genetics––is notantithetical to at least one environmentaltheory––anomie/strain theory.

y Behavior Genetics

I suggest that it is time for mainstream criminol-ogy to at least pull back its blinders and peek atwhat behavior genetics has to offer. Very few crim-inologists have taken biology beyond an introduc-tory class, and even fewer believe that geneticfactors are important in explaining criminalbehavior, despite the overviews of biosocial theo-ries that have been published in the past decade.The lack of human genetic knowledge is probablynot as important as ideology in keeping our blind-ers on. To acknowledge that genes may play a role in criminality has been virtually taboo fordecades, and many criminologists still suffer from“biophobia,” to use Ellis’s (1996a) expression.

Behavior genetics, a branch of quantitativegenetics, is not so much a “biological” discipline asit is a biologically-friendly environmental disci-pline. Although the discipline’s main focus is tounderstand genetic influences on human behavior,traits, and abilities, behavior geneticists are awarethat this cannot be accomplished without under-standing the complementary role of the environ-ment. Behavioral genetic studies have typicallyfound that the environment accounts for morevariance than does genetics in the most humantraits, behaviors, and abilities. Thus, an often unap-preciated aspect of behavior genetics is that it tellsus as much, or sometimes more, about environ-mental effects as it does about genetic effects.

Behavior genetics research designs can alsoaddress and rectify a major problem with whathas been called the standard social science model(SSSM) of socialization (Tooby and Cosmides,1992). The problem is that the SSSM can neverdetermine if any observed effects are primarilygenetically or environmentally driven, i.e., whetherparent/child or sibling/sibling similarities arecaused primarily by shared genes or shared envi-ronments. This has led some socializationresearchers to dismiss the entire SSSM of social-ization as essentially useless.

We can only disentangle genetic from envi-ronmental effects if we can hold one constant.Behavior genetics does this using twin and adop-tion studies. The twin method compares pairs ofindividuals reared together (e.g., monozygotictwins) with a known degree of genetic related-ness with other pairs of reared-together individ-uals with a different degree of genetic relatedness(e.g., dizygotic twins), which holds environmentsconstant and allows genes to vary. The adoptionmethod examines genetically related individualsreared in different homes (or genetically unre-lated individuals reared in the same home),which holds genes constant and allows environ-ments to vary. In other words, the effects ofshared environment can be determined by study-ing genetically unrelated individuals raised in thesame environment, and the effects of hereditycan be gauged by the phenotypical similarities ofgenetically related individuals reared apart.

Until molecular genetics identifies specificgenes underlying specific traits, genetic effectsmust be inferred rather than directly demon-strated. Genetic effects are inferred by calculatinga trait’s heritability (h2). Heritability coefficientsrange between 0 and 1 and estimate the extent towhich variance in a trait in a population is attrib-utable to genes. Heritability is based on theassumption that if genes affect a trait, the moregenetically related two individuals are, the moresimilar they will be on that trait. If genes do notaffect a trait, it would be logically and empiricallyimpossible to calculate a heritability coefficientfor that trait significantly greater than zero.

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Heritability estimates fluctuate among differ-ent populations and within the same populationas they experience different environments.Knowing what percentage of variance in a trait isattributable to genetic factors does not set limitson creating other environments that may influ-ence the trait. A large heritability coefficientinforms us that the present environment at thepresent time has minimal effect (accounts forlittle variance) on the trait; it does not tell us whatother environments may have appreciably greatereffects on the trait. Heritability tells us what isaffecting trait variance; it does not tell us what canaffect it. Moreover, the heritability of a traitinforms us only about how much of the variancein a trait is attributable to genes, not how muchthe trait is attributable to genetic influences.

A related misunderstanding about heritabil-ity is the assumption that it carves nature neatly atthe joint because it apportions traits into “genetic”and “environmental” components (e.g., 60%genetic, therefore, 40% environmental). Althoughheritability estimates statistically transform genesand environments into components, they are notseparable ingredients in the real world. Genes andenvironments have the same relationship to phe-notypic attributes as hydrogen and oxygen have towater and length and width have to area. It takestwice as many hydrogen atoms as oxygen atoms tomake a molecule of water and an area’s length maybe twice its width, but in terms of the wholes theydescribe, the quantitative differences are meaning-less. Without their complements, there would beno water or area, only a gas and a straight line.Likewise, genes and environments in isolationmake no sense in terms of the phenotypic wholesthey describe. There is no nature versus nurture,there is only nature via nurture.

In addition to apportioning variance intogenetic and environmental components, themethods of behavior genetics yield a further ben-efit to social science in that they allow researchersto break down environmental variance intoshared and nonshared components. Shared envi-ronments are environments of rearing that serveto make siblings alike (e.g., parental SES, religion,

neighborhood, intactness of home), and non-shared environments are microenvironmentsunique to each sibling (birth order, differentialtreatment, different peers, etc.) that serve to makethem different. A cascade of behavior geneticresearch has shown unequivocally that althoughshared rearing effects are real, albeit modest, dur-ing childhood and adolescence, they almost com-pletely disappear in adulthood.

y Agency and Gene/Environment Correlation

Social scientists are increasingly acknowledgingthat people’s choices are not passive responses tosocial and cultural situations, and that thesechoices are to some degree autonomous. Manysocial scientists have gravitated toward the con-cept of agency in response to the “oversocialized”conception of human nature that has for so longdominated the social sciences. Agency simplymeans that as individuals strive for autonomy,they affect their environments just as surely asthey are affected by them.

Transformations of self and environmentare achieved by the actions and interactions ofhuman subjects propelled by the subjectivemeanings that different people assign to similarsituations. This concept of agency, which hasmuch to do with our wishes, goals, and desires, ishighly congenial to behavior genetics, which hasalways emphasized that people make their envi-ronments. That is, people’s unique genotypes willlargely determine what aspects of the social envi-ronment will be salient to them. This is a positionconsiderably more respectful of human dignity(and more scientifically defensible) than is theimage of human development that views it aslittle more than a process of class-, race-, age-, orgender-based adjustment to structural and cul-tural demands made on us. The concept ofagency pulls us away from thinking of socializa-tion as a mechanistic parent-to-child process andprovides us with the skeleton of reciprocal effectsthinking. Behavior genetics goes a step further toput the flesh on the bones of this thinking.

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The concept of gene/environment (G/E)correlation is philosophically related to the con-cept of agency, but goes beyond it to provide anunderstanding of the underlying mechanismsthat constitute the basis for the subjective mean-ings agency theorists articulate. G/E correlationessentially means that genotypes and the envi-ronments they encounter are not random withrespect to one another, and that individuals areactive shapers of their lives. The concept enablesus to conceptualize the indirect way that genesexert their influence to help to determine theeffective environment of the developing individ-ual. Behavior geneticists differentiate betweenpassive, reactive, and active G/E correlation.

Children raised by their biological parentsexperience passive G/E correlation by virtue ofbeing provided with genes that underlie certaintraits and a home environment favorable fortheir expression. A child born to highly intelli-gent parents, for example, typically receives genesconducive to high intelligence and a home inwhich intellectual activity is modeled and rein-forced. The synergistic effects of correlated genesand environment will make for the unfolding ofthe child’s intellectual abilities almost indepen-dently (passively) of what the child does. Thismeans that the child has simply been exposed tothe environment and has not been instrumentalin forming it, not that the child does not activelyengage it. On the downside, children born to lowIQ, impulsive, or bad-tempered parents receiveboth genes and an environment developmentallybiasing them in the same direction. The influenceof passive G/E correlation declines dramaticallyfrom infancy to adolescence as the scope of envi-ronmental interaction widens and the person isconfronted with and engages a wider variety ofother people and behavioral options.

Reactive G/E correlation picks up the devel-opmental trajectory as children grow older andare exposed to an increasing number of peopleand situations in their environments and beginto respond more actively to them. Children bringtheir developing phenotypical characteristics andabilities with them to the interpersonal situations

they encounter that increase or decrease theprobability of evoking certain kinds of responsesfrom others. A bad-tempered child will evoke lesssolicitude than will a good-natured child, and ahyperactive, moody, and mischievous child willevoke less benign responses from others than willone who is pleasant and well behaved. Likewise, achild who shows enthusiasm and ability forschool work evokes better treatment from teach-ers and will be afforded greater opportunities foradvancement than will a child who shows littleenthusiasm and ability for school work.

The important lesson of reactive G/E corre-lation is that the behavior of others toward thechild is as much a function of the child’s evoca-tive behavior as it is of the interaction style ofthose who respond to it. Socialization is notsomething that others simply do to children; it isa reciprocal process that children and their care-takers engage in together. Some children may bedifficult to control for even the most patient andloving parents. Some parents may abuse difficultchildren in their efforts to make them conform,and other parents may just give up trying tosocialize their children. Both abusive and permis-sive responses to difficult children are likely toexacerbate their antisocial tendencies and drivethem to seek others more accepting of their ten-dencies, presumably because the others towardwhom they gravitate harbor such tendencies.Groups of individuals with similar tendenciesprovide positive reinforcement for each otherand provide ample opportunities to exercisethese tendencies. It is in this way that the feed-back nature of reactive G/E correlation amplifiesdifferences among phenotypes.

The seeking out of environments in whichone feels accepted and psychologically comfort-able is referred to as active G/E correlation.Large-scale twin studies provide striking evi-dence that genes play a very important role in“niche-picking.” A number of studies havereported that the similarity in intelligence, per-sonalities, attitudes, interests, and constructedenvironments of monozygotic (MZ) twins isessentially unaffected by whether they were

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Section 8 � Behavior Genetics and Anomie/Strain Theory 291

reared together or apart. That is, MZ twins rearedapart construct their environments and ordertheir lives about as similarly as they would havehad they been raised together, and this similarityis considerably greater than is the similaritybetween dizygotic (DZ) twins reared together.

Although such findings are robust, theyappear counterintuitive to those who believe thatthe rearing environment mostly determines lifeoutcomes. However, they make perfect sense tobehavior geneticists, for whom it would be coun-terintuitive for people to either accept or to seek out environments incompatible with theirgenetic dispositions. Because MZ twins share100% of their genes and DZ twins share, on aver-age, only 50%, it makes sense that MZ twinswould order their lives more similarly thanwould DZ twins. People with genes facilitative ofdifferent temperaments, traits, and abilities willseek out environments that mesh well with them(genes and environments will covary positively).It is no surprise to behavior geneticists thatwithin the range of cultural possibilities and con-straints, our genes set us on a developmental tra-jectory that will largely determine what featuresof the social world will be meaningful andrewarding to us, and what features will not.

y Behavior Genetics and Antisocial Behavior

I must strongly emphasize that there isabsolutely nothing in the concept of G/E corre-lation that can in any way be construed as sup-porting the notion of congenital criminality.Genes are self-replicating slices of DNA thatcode for proteins, which code for hormonal andenzymatic processes. There is no mysteriouscryptography by which genes code for certainkinds of brains, which in turn code for differentkinds of behaviors. Genes do not code for feel-ings or emotions either; what they do is makeus differentially sensitive to environmentalcues and modulate our responses to them.Genes always exert whatever influence they haveon behavior in an environmental context.

Although there can be no gene(s) “for” crime,there are genes that, via a number of neurohor-monal routes, lead to the development of differ-ent traits and characteristics that may increasethe probability of criminal behavior in someenvironments and in some situations.

A recent behavior genetic study illustratesthe role of G/E correlation for antisocial behaviorin late childhood/early adolescence (O’Connor et al., 1998). A number of adopted children wereclassified as either being or not being at geneticrisk for antisocial behavior on the basis of theirbiological mothers’ self-reported antisocial behav-ior collected prior to the birth of their children. Itwas found that from ages 7 to 12, children atgenetic risk for antisocial behavior consistentlyreceived more negative parenting from theiradoptive parents than did children not at geneticrisk. This effect was interpreted as reactive G/Ecorrelation in that children’s poor behavior wasseen as evoking negative parenting. O’Connor et al. (1998) did, however, find an environmen-tally mediated parental effect not attributable toreactive G/E correlation.

Another adoption study of antisocial behav-ior focused on G/E interaction (the differentialeffects of similar environments on differentgenotypes). Cadoret et al. (1995) examined theantisocial history of adopted children separatedat birth from biological mothers with verifiedantisocial histories, compared with otheradoptees with biological mothers with no knownhistory of antisocial behavior. It was found thatadverse adoptive home environments (divorce/separation, substance abuse, neglect/abuse, mar-ital discord) led to significant increases in antiso-cial behavior for adoptees at genetic risk, but notfor adoptees without genetic risk. Thus, bothgenes and environments operating in tandem(interacting) were required to produce signifi-cant antisocial behavior, whereas neither seemedpowerful enough in this study to produce sucheffects independent of the other.

Antisocial behaviors, especially adolescentantisocial behaviors, are an interestingexception to the modest shared environmental

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influences typically discovered for most humancharacteristics. Shared group influences reflectthe socializing influences of peer groups,although such influences cannot be consideredapart from the tendency of similar people tobefriend one another. The shifting pattern ofgenetic and environmental effects is readilyseen in studies of juvenile and adult offending.

Genetic factors do not have similar explana-tory power across all environments and across alldevelopmental periods. Few things point to thevital importance of the environment to geneexpression than the fact that heritability coeffi-cients are always higher in advantaged than indisadvantaged environments. Just as a rose willexpress its fullest genetic potential planted in anEnglish garden and wither when planted in theNevada desert, human beings will realize theirgenetic potential to the fullest when reared inpositive environments and fall short of doing sowhen reared in negative environments. High her-itabilities for positive traits index how well asociety is doing in equalizing environmentalopportunities.

Differential genetic effects also apply todelinquent and criminal behaviors. In environ-ments where resistance to crime is low, very littlevariance in criminal behavior will be attributableto genes; in environments where resistance tocrime is high, the genetic contribution will behigh. Venables (1987), for example, found that(low) tonic heart rate was a significant predictorof antisocial behavior among high SES children,but not among low SES children. Similarly, Walsh(1992) found that cognitive imbalance (as mea-sured by verbal/performance IQ discrepancyscores) significantly predicted violent delin-quency in advantaged environments, but not indisadvantaged environments. This does notmean these studies found that low tonic heartrate or cognitive imbalance was more prevalentin advantaged environments. On the contrary,what it means is that environmental causes tendto overwhelm putative genetic causes in disad-vantaged environments.

The environmental complexities illustratedby studies in this section make criminal/antisocialbehavior especially appealing for behaviorgenetic analysis of environmental influence.

y Traits Linked to Middle-Class Success

In the following section, I briefly review the liter-ature on the two major individual-level factors(temperament and intelligence) stressed byAgnew [in his general strain theory] (1992, 1997)as they relate to achieving middle-class success.Agnew remarks on a number of occasions thattemperament and intelligence bear a strongrelationship to problem-solving skills, and thatthe lower classes feel strain most acutely(1997:111–114). He never tries to make the con-nection between SES and these correlates ofproblem solving, although he is more than will-ing to state that such traits are a function of bothbiological and social factors. After the discussionof temperament and intelligence, I will attemptto show how other biosocial variables can be use-fully integrated with Agnew’s (1997) recently for-mulated developmental version of GST.

Temperament

Temperament refers to an individual’s habit-ual mode of emotionally responding to stimuli.Temperamental style is identifiable very early inlife, and it tends to be stable across the life course.Variance in temperamental measures is largely afunction of heritable variation in central andautonomic nervous system arousal. Heritabilitycoefficients for the various components of temp-erament range from about .40 to .80. Tempera-mental differences are largely responsible formaking children differentially responsive tosocialization. The unresponsiveness of a badtempered (sour, unresponsive, quick to anger)child is exacerbated by the fact that tempera-ments of children and their parents are typicallypositively correlated. Parents of children with

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Section 8 � Behavior Genetics and Anomie/Strain Theory 293

difficult temperaments tend to be inconsistentdisciplinarians, irritable, impatient, and unstable,which makes them unable or unwilling to copeconstructively with their children. Their childrenare thus burdened with both a genetic and anenvironmental liability. A cascade of evidenceshows that children with difficult temperamentsevoke negative responses from parents, teachers,and peers, and that these children find accep-tance only in association with peers with similardispositions (Moffitt, 1996).

Various physiological measures confirm thatdisinhibited temperament is associated with central nervous system (CNS) and peripheralnervous system (PNS) underarousal (suboptimallevels of arousal under normal environmentalconditions). Individuals differ in the level ofenvironmental stimulation they find optimalbecause of variation in the CNS’s reticular acti-vating system (a finger-size cluster of cellsextending from the spinal cord that monitorsincoming stimuli for processing by the cerebralcortex), and the PNS’s autonomic nervous sys-tem. What is optimal for most of us will be stress-ful for some and boring for others. Suboptimallyaroused people are easily bored and continuallyseek to boost stimuli to more comfortable levels.This search for amplified pleasure and excite-ment often leads the underaroused person intoconflict with the law. A number of studies haveshown that relative to the general population,criminals, especially those with the most seriouscriminal records, are chronically underaroused asdetermined by EEG brain wave patterns, restingheart rate and skin conductance, and histories ofhyperactivity and attention deficit disorders.Individuals who are chronically bored and con-tinually seeking intense stimulation are not likelyto apply themselves to school or endear them-selves to employers. One study found that vari-ous measures of reticular and autonomicnervous system arousal taken at age 15 correctlyclassified 74.7% of subjects as “criminal” or“noncriminal” at age 24 after controlling for anumber of social factors (Raine et al., 1990).

Temperament provides the foundations forpersonality, which emerges from its interactionwith the environment. Conscientiousness, one ofthe “big five” factors of personality, is a trait thatis particularly important to success in the workforce and is thus important to anomie/straintheory. Conscientiousness is a continuous traitranging from well organized, disciplined, reli-able, responsible, and scrupulous at one end ofthe continuum, to disorganized, careless, unreli-able, irresponsible, and unscrupulous at theother. As we might expect, variance in conscien-tiousness is heritable. An analysis of 21 behaviorgenetic studies of conscientiousness found amedian heritability of .66. In short, individualswith temperaments biasing them in the directionof being lacking in conscientiousness typically donot possess the personal qualities needed toapply themselves to the often long and arduoustask of achieving legitimate occupational success.They will become “innovators’ or “retreatists,”not because a reified social structure has deniedthem access to the race, but because they find therace intolerably boring and busy themselves withmore “exciting” pursuits instead.

Intelligence

Intelligence, as operationalized by IQ tests, isanother obvious determinant of both occupa-tional success and coping strategy, but also onethat is conspicuously absent in sociological dis-cussions of social status. As Lee Ellis has opined:“Someday historians of social science will beastounded to find the word intelligence is usuallynot even mentioned in late-twentieth-centurytext books on social stratification” (1996b:28).

Few scientists who study intelligence seri-ously doubt the importance of genes as well asthe environment to explaining IQ variation.Twin studies, adoption studies, position elec-tron tomography and magnetic resonanceimage scan studies, and even molecular geneticstudies all point to substantial genetic effects onintelligence. Contrary to the claims of many

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social scientists, the National Academy ofSciences and the American PsychologicalAssociation’s Task Force on Intelligence haveconcluded that there is no empirical evidence toindicate that IQ tests are biased against anyracial/ethnic group or social class.

The litmus test for any assessment tool is itscriterion-related validity—its ability to predictoutcomes. An examination of 11 meta-analysesof the relationship between IQ and occupationalsuccess found that IQ predicted success betterthan did any other variable in most occupations,particularly in higher status occupations, andthat it predicted equally well for all races/ethnicgroups. Intelligence is particularly important inopen and technologically advanced societies.Although there is no such thing as a totally opensociety, unlike the rigid caste-like societies of thepast. In the rigid and aristocratic caste societies ofthe past, genes played almost no role in deter-mining social class. They play an increasinglyimportant role in more modern and egalitariansocieties, however. Genes, and the individual dif-ferences they underlie, become important indetermining SES in roughly direct proportion toequalization of environments. Although this mayseem paradoxical at first blush, it is a basic prin-ciple of genetics: The more homogeneous (orequal) the environment, the greater the heritabil-ity of a trait; the more heterogeneous (unequal)the environment, the lower the heritability of atrait. High heritability coefficients for sociallyimportant traits tell us that the society is doing agood job of equalizing the environment.

The degree of occupational mobility in theU.S. labor force can be gauged by a major study’sfinding that 48% of sons of upper white-collarstatus fathers had lower status occupations thandid their fathers, with 17% falling all the way to“lower manual” status, and that 51% of sons oflower manual status fathers achieved higher sta-tus, with 22.5% achieving “upper white-collar”status (Hurst, 1995: 270). Given this degree ofupward/downward social mobility, and given thedegree to which IQ predicts it equally for all races

and social classes, it is difficult to maintain thatany group is systematically denied access to legit-imate opportunities to attain middle-class status.

Social scientists are more prone to attributeIQ level to SES level rather than the other wayaround. A large number of studies have foundthe correlation between parental SES andchildren’s IQ to be within the .30 to .40 range(which is predictable from polygenetic transmis-sion models). However, the correlation betweenindividuals’ IQ and their attained adult SES is inthe .50 to .70 range. As Jensen remarks: “If SESwere the cause of IQ [rather than the other wayaround], the correlations between adults’ IQ andtheir attained SES would not be markedly higherthan the correlation between children’s IQ andtheir parents’ SES” (1998: 491).

If differential IQ predicts differential adultSES, and if the lack of success leads to a mode ofadaptation that includes criminal activity, IQmust be a predictor of criminal behavior. Whenevaluating the relatively small (eight IQ points)difference said to separate criminals and non-criminals, we must remember that researchersdo not typically separate what Moffitt (1993)calls adolescent-limited (AL) and “life-course-persistent” (LCP) offenders. As statistically nor-mal individuals responding to the contingenciesof their environments, we would not expect ALoffenders to be significantly different from non-offenders on IQ, and they are not. Moffitt (1996)reports a one-point mean IQ deficit between ALoffenders and nonoffenders, but a 17-pointdeficit between LCP offenders and nonoffend-ers, and Stattin and Klackenberg-Larsson(1993) found a mean deficit of 4 points betweennonoffenders and “sporadic” offenders and a10-point difference between nonoffenders and“frequent” offenders. Aggregating temporaryand persistent offenders creates the erroneousperception that IQ has minimal impact on anti-social behavior.

Although these studies separated tempo-rary and persistent offenders, they did not sepa-rate IQ subtest scores. This also leads to an

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Section 8 � Behavior Genetics and Anomie/Strain Theory 295

underestimation of the effects of IQ by poolingverbal IQ (VIQ), which uniformly shows a sig-nificant difference between offenders andnonoffenders, and performance IQ (PIQ),which typically does not. The most serious andpersistent criminal offenders tend to have a PIQscore exceeding their VIQ score by about 12points. Low verbal IQ (about one standard devi-ation below the mean) indexes poor abstractreasoning, poor judgment, poor school perfor-mance, impulsiveness, low empathy, and pre-sent orientedness. None of these traits isconducive to occupational success, but it is con-ducive to antisocial behavior, especially if com-bined with a disinhibited temperament.

y Agnew’s Developmental GST and Other Biosocial Processes

Agnew’s (1997) developmental strain theoryattempts to explain why antisocial behaviorpeaks during adolescence, focusing on increasesin negative relationships with others during thisperiod and the tendency to cope with the result-ing strain through delinquency. Thus, a numberof situations experienced as aversive arehypothesized to account for the age effect oncrime and delinquency. As other theorists havepointed out, however, the age effect tends toremain robust, controlling for a number ofdemographic and situational variables, indicat-ing that none of the various social correlates ofage predict crime as well as age. For instance,Udry’s (1990) sociological model found that ageremained the strongest predictor of “problembehavior,” controlling for a number of otherindependent variables. However, in his biosocialmodel that included measures of testosterone(T) and sex hormone binding globulin (SHBG),age dropped out of the equation, promptinghim to suggest that age is a proxy for the hor-monal changes of puberty.

During puberty, male T-levels increase 10-to 20-fold and SHBG is halved. Although the heritability of base-level T is around .60, the

environmental effects on this hormone are wellknown. It is plain that T/SHBG ratios alone can-not explain the dramatic increase in antisocialbehavior during this period, nor can they explainthe fairly rapid decline in such behavior in thelate teens and early twenties, because the declinedoes not correspond with a similar decline inT. Whatever the environmental or hormonalcorrelates of the increase and decline in adoles-cent antisocial behavior are, they are necessarilymediated by the brain.

Recent MRI studies of brain developmentconfirm that the prefrontal cortex (PFC) is thelast part of the human brain to fully mature. ThePFC is the part of the brain that serves variousexecutive functions, such as modulating emo-tions from the more primitive limbic system andmaking reasoned judgments. The PFC undergoesan intense prepubescent period of synaptogene-sis and a period of pruning of excess synapsesduring adolescence. This process of synapticelimination may be part of the reason manyyoung persons find it difficult to accurately gaugethe meanings and intentions of others and toexperience more stimuli as aversive.

In addition to the synaptic pruning process,the adolescent’s PFC is less completely myeli-nated than is the adult PFC. Myelination isimportant to the speed and conductive efficiencyof neurotransmission. The fact that many syn-dromes associated with delinquency, such asoppositional disorder and conduct disorder/socialized type, first appear during this periodmay reflect a brain that is sometimes develop-mentally not up to the task of dealing rationallywith the strains of adolescence. For instance,magnetic resonance imaging studies have shownthat only the emotional limbic system is activatedfor many adolescents when shown photographsof frightened people, whereas both the limbicsystem and the PFC show activity among matureadults. This is another indication that the PFC is probably not performing its reasoning andemotion-modulating duties efficiently for at leastsome adolescents.

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The increase in behavior-activating hor-mones coupled with an immature brain reflecttwo biological processes temporarily on con-flicting trajectories that may both generate andexacerbate the strains of adolescence. Thisconjecture is supported by studies indicatingthat boys entering puberty early throw signifi-cantly more temper tantrums than do latermaturing boys, and that early-maturing girlsengage in significantly more problem behav-iors than do later-maturing girls. Also consis-tent with this is the finding that T-levelspredict future problem behavior only for early-pubertal-onset boys, again implying that theimmature adolescent brain may facilitate a ten-dency to assign faulty attributions superim-posed on an unfamiliar and diffuse state ofphysiological arousal.

Many other neurohormonal processes havethe potential to assist anomie/strain theorists tounderstand strain across the life span, particu-larly during the teenage years. These include theshifting ratios of behavior-facilitating dopamineand the behavior-moderating serotonin duringadolescence. The deaminating enzyme mono-amine oxydase, which removes excess neuro-transmitters at the synapse, is at its lowest duringthis period. These neurochemical fluctuationssuggest that adolescence may be a period inwhich the brain is particularly sensitive torewards and relatively insensitive to punish-ment, a combination facilitative of risk-takingand sensation-seeking behaviors.

It has long been known that humans andother animals in a low serotonin state are proneto violence, impulsiveness, and risk taking,especially if combined with high T. However,none of this means that teenagers are at themercy of their biology, or that the environmenthas little impact. Serotonin levels, like T-levels,largely reflect environmental influences, andthe lower average levels of serotonin during

adolescence are as likely to be the effect of theincreased strains of this period as they are to be the cause (Bernhardt, 1997). The stresses ofadolescence outlined in Agnew’s (1997) theoryprompt the release of corticosteroids, which inturn suppress serotonin receptors. Thus, thevicissitudes of biology and social situationsduring this period of life combine in synergis-tic fashion to become both the producers andthe products of strain. Further exploration ofthese interesting processes is beyond the scopeof this paper.

y Conclusion

I have attempted to show how behavior geneticscan complement, extend, and add coherence tocriminological theory, using one of its mostrevered and long-lived theories as an illustration.Perhaps all social scientists would acknowledgein the abstract that human behavior is the resultof complex interactions between genes and envi-ronments, but having done so, most will con-tinue to neglect the genetic 50% of the humanbehavioral equation in their thinking andresearch. As I have repeatedly emphasized,behavior genetics is not a “biological” perspectiveof human behavior. It is a biosocial perspectivethat takes seriously the proposition that allhuman traits, abilities, and behaviors are theresult of the interplay of genetics and environ-ments, and it is the only perspective with theresearch tools to untangle their effects. Given theexponential increase in knowledge in geneticsover the past decades, criminologists cannot con-tinue to reject insights from behavior genetics ifthe discipline is to remain scientifically viable.The history of science tells us that cross-discipli-nary fertilization by a more mature science has,without exception, proven to be of immensevalue to the immature science. Criminology willbe no exception.

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Section 8 � Behavior Genetics and Anomie/Strain Theory 297

y ReferencesAgnew, R. (1992). Foundations for a general strain theory of

crime and delinquency. Criminology, 30, 47–87.Agnew, R. (1997). Stability and change in crime over the life-

course: A strain theory explanation. In T. Thornberry(Ed.), Developmental theories of crime and delinquency.New Brunswick, NJ: Transaction.

Bernhardt, P. (1997). Influences of serotonin and testos-terone in aggression and dominance: Convergence withsocial psychology. Current Directions in PsychologicalScience, 6, 44–48.

Cadoret, R., Yates, W., Troughton, E., Woodworth, G., &Stewart, M. (1995). Genetic-environmental interactionin the genesis of aggressivity and conduct disorders.Archives of General Psychiatry, 52, 916–924.

Durkheim, E. (1982). The rules of sociological method. NewYork: Macmillan.

Ellis, L. (1996a). A discipline in peril: Sociology’s futurehinges on curing its biophobia. American Sociologist,27, 21–41.

Ellis, L. (1996b). Arousal theory and the religiosity-criminalityrelationship. In P. Cordella & L. Siegel (Eds.), Readings incontemporary criminological theory. Boston: NortheasternUniversity Press.

Hurst, C. (1995). Social inequality: Forms, causes, and conse-quences. Boston: Allyn & Bacon.

Jensen, A. (1998). The g factor. Westport, CT: Praeger.Moffitt, T. (1993). Adolescent-limited and life-course persis-

tent antisocial behavior: A developmental taxonomy.Psychological Review, 100, 674–701.

Moffitt, T. (1996). The neuropsychology of conduct disorder.In P. Cordella & L. Siegal (Eds.), Readings in contemporary

criminological theory. Boston: Northeastern UniversityPress.

O’Connor, T., Deater-Deckard, K., Fulker, D., Rutter, M., &Plomin, R. (1998). Genotype-environment correlationsin late childhood and early adolescence: Antisocialbehavioral problems and coercive parenting.Developmental Psychology, 34, 970–981.

Raine, A., Venables, P., & Williams, M. (1990). Relationshipsbetween central and autonomic measures of arousal atage 15 years and criminality at age 24 years. Archives ofGeneral Psychiatry, 47, 1003–1007.

Stattin, H., & Klackenberg-Larsson, I. (1993). Early languageand intelligence development and their relationship tofuture criminal behavior. Journal of Abnormal Psychology,102, 369–378.

Tooby, J., & Cosmides, L. (1992). The psychological founda-tions of culture. In J. Barkow, L. Cosmides, & J. Tooby(Eds.), The adapted mind: Evolutionary psychology andthe generation of culture (pp. 19–136). New York: OxfordUniversity Press.

Udry, J. R. (1990). Biosocial models of adolescent problembehaviors. Social Biology, 37, 1–10.

Udry, J. R. (1995). Sociology and biology: What biology dosociologists need to know? Social Forces, 73, 1267–1278.

Venables, P. (1987). Autonomic nervous system factors incriminal behavior. In S. Mednick, T. Moffitt, & S. Stack(Eds.), The causes of crime: New biological approaches(pp. 110–136). Cambridge: University of CambridgePress.

Walsh, A. (1992). Genetic and environmental explanationsof violent juvenile delinquency in advantaged and dis-advantaged environments. Aggressive Behavior, 18,187–199.

D I S C U S S I O N Q U E S T I O N S

1. How does the concept of G/E correlation highlight the role of human agency in development?

2. Why is talking about the role of intelligence in occupational success (and crime) controversial?

3. Outline the ways in which behavior genetics has thrown light on the anomie/strain tradition.

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READING

Neuroimaging Studies of Aggressive and Violent Behavior

Current Findings and Implications for Criminology and Criminal Justice

Jana L. Bufkin and Vickie R. Luttrell

This article reviews neuroimaging studies with the goal of showing how neuroscience can advancecriminological theories and inform criminal justice practice. The availability of new functional andstructural neuroimaging techniques has allowed researchers to localize brain areas that may be dys-functional in offenders who are aggressive and violent. Bufkin and Luttrell’s review of 17 neu-roimaging studies reveals that the areas associated with aggressive and/or violent behavioralhistories, particularly impulsive acts, are located in the prefrontal cortex and the medial temporalregions. These findings are explained in the context of negative emotion regulation, thus support-ing many studies outside of neuroscience that link criminality to low self-control and negativeemotionality. Bufkin and Luttrell provide suggestions concerning how such findings may affectfuture theoretical criminology, crime prevention efforts, and treatment in criminal justice.

298 INTRODUCTION TO CRIMINOLOGY

SOURCE: Bufkin, J. L., & Luttrell, V. R. (2005). Neuroimaging studies of aggressive and violent behavior: Current findings andimplications for criminology and criminal justice. Trauma, Violence, & Abuse, 6(2), 176–191. Reprinted with permission of SagePublications, Inc.

Aggressive and/or violent behaviors persist assignificant social problems. In response, a sub-stantial amount of research has been conductedto determine the roots of such behavior. Casestudies of patients with neurological disorders orthose who have suffered traumatic brain injuryprovide provocative insights into which brainregions, when damaged, might predispose toirresponsible, violent behavior. Psychophy-siological and neuropsychological assessmentshave also demonstrated that violent offendershave lower brain functioning than controls,including lower verbal ability and diminishedexecutive functioning. However, until recently ithas been impossible to determine which brainareas in particular may be dysfunctional in

violent individuals. With the availability of newfunctional and structural neuroimaging tech-niques, such as single-photon emission com-puted tomography (SPECT), positron emissiontomography (PET), magnetic resonance imaging(MRI) and functional MRI (fMRI), it is now pos-sible to examine regional brain dysfunction witha higher sensitivity and accuracy than was possi-ble with previous techniques. This newfoundability to view the brain “in action” has broad-ened our understanding of the neural circuitrythat underlies emotional regulation and affiliatedbehaviors. In particular, evidence suggests thatindividuals who are vulnerable to faulty regula-tion of negative emotion may be at increased riskfor aggressive and/or violent behavior.

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Section 8 � Neuroimaging Studies of Aggressive and Violent Behavior 299

In this review, we evaluate the proposed linkbetween faulty emotion regulation and aggres-sive or violent behavior. We define aggression asany threatening or physically assaultive behaviordirected at persons or the environment. Violencerefers to behaviors that inflict physical harm inviolation of social norms. Specifically, we (a) dis-cuss briefly the neurobiology of emotion regula-tion and how disruptions in the neural circuitryunderlying emotion regulation might predisposeto impulsive aggression and violence; (b) sum-marize the results of modern neuroimagingstudies that have directly assessed brain func-tioning and/or structure in aggressive, violent,and/or antisocial samples and evaluate the consistency of these findings in the context ofnegative emotion regulation; and (c) discuss the-oretical and practical implications for criminol-ogy and criminal justice.

y Emotion Regulation andTheoretical Links to ImpulsiveAggression and Violence

Emotion is regulated by a complex neural circuitthat involves several cortical areas, including theprefrontal cortex, the anterior cingulate cortex(ACC), the posterior right hemisphere, the insu-lar cortex, as well as several subcortical struc-tures, such as the amygdala, hippocampus, andthalamus. These cortical and subcortical areas areintricately and extensively interconnected. In thisarticle, we focus on three key elements of thisneural circuitry: the prefrontal cortex, the ACC,and the amygdala.

The prefrontal cortex is a histologicallyheterogeneous region of the brain and has sev-eral (somewhat) functionally distinct sectors,including the ventromedial cortex and theorbitofrontal cortex (OFC). Damage to the ven-tromedial cortex and its behavioral affiliationshave been assessed in case studies of individualswho experienced traumatic brain injury, eitherduring childhood or adulthood, and in large,systematic studies on cohorts of war veteranswith head injury.

Studies have found that patients with early-onset ventromedial lesions experience an insen-sitivity to future consequences, an inability tomodify so-called risky behaviors even whenmore advantageous options are presented, anddefective autonomic responses to punishmentcontingencies. Studies have also demonstratedthat patients with adult-onset ventromedialdamage show defects in real-life decision mak-ing, are oblivious to the future consequences oftheir actions, seem to be guided by immediateprospects only, and fail to respond autonomicallyto anticipated negative future outcomes.

The OFC, also a part of the prefrontal cir-cuit, receives highly processed sensory informa-tion concerning a person’s environmentalexperiences. The OFC is hypothesized to play arole in mediating behavior based on social con-text and appears to play a role in the perceptionof social signals, in particular, facial expressionsof anger. Blair et al. (1999), using PET scans,assessed 13 male volunteers as they viewed staticimages of human faces expressing varyingdegrees of anger. They found that increasing theintensity of angry facial expressions was associ-ated with enhanced activity in participants’ OFCand the ACC. Dougherty et al. (1999) used func-tional neuroimaging and symptom provocationtechniques to study the neurobiology of inducedanger states and found that imaginal anger wasassociated with enhanced activation of the leftOFC, right ACC (affective division), and bilateralanterior temporal regions. Also using imaginalscenarios, Pietrini et al. (2000) found that func-tional deactivation of OFC areas was strongestwhen participants were instructed to expressunrestrained aggression toward assailants ratherthan when they tried to inhibit this imaginalaggression. Taken together, these lines of evi-dence support the suggestion that heightenedactivity in the left OFC may prevent a behavioralresponse to induced anger.

Based on these findings, and consistent withfearlessness theories of human aggression, a log-ical prediction is that OFC and ACC activity inresponse to provocation may be attenuated in

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certain individuals, predisposing them to aggres-sion and violence. Consistent with this predic-tion, patients with OFC damage tend to exhibitpoor impulse control, aggressive outbursts, ver-bal lewdness, and a lack of interpersonal sensitiv-ity, which may increase the probability ofsporadic so-called crimes of passion and encoun-ters with the legal system. In contrast, evidencesuggests that the ACC plays a role in processingthe affective aspects of painful stimuli, such asthe perceived unpleasantness that accompaniesactual or potential tissue damage.

In addition to the prefrontal cortex and theACC, another hypothesized neural component ofemotion regulation is the amygdala, a subcorticalstructure, which is located on the medial marginof the temporal lobes. Similar to the OFC, theamygdala appears to play a role in extractingemotional content from environmental stimuliand may also play a role in individuals’ ability toregulate negative emotion. However, neuroimag-ing studies have found that the amygdala is acti-vated in response to cues that connote threat,such as facial expressions of fear (instead ofanger), and that increasing the intensity of fear-ful facial expressions is associated with anincreased activation of the left amygdala.

Davidson, Putnam, et al. (2000) suggestedthat individuals can typically regulate their nega-tive affect and can also profit from restraint-producing cues in their environment, such asothers’ facial expressions of fear or anger.Information about behaviors that connote threat(e.g., hostile stares, threatening words, or lungingpostures) is conveyed to the amygdala, which thenprojects to other limbic structures, and it is therethat information about social context derivedfrom OFC projections is integrated with one’s cur-rent perceptions. The OFC, through its connec-tions with other prefrontal sectors and with theamygdala, plays an important role in inhibitingimpulsive outbursts because prefrontal activationsthat occur during anger arousal constrain theimpulsive expression of emotional behavior.

Davidson, Putnam, et al. (2000) also pro-posed that dysfunctions in one or more of these

regions and/or in the interconnections amongthem may be associated with faulty regulation ofnegative emotion and an increased propensityfor impulsive aggression and violence. First,people with prefrontal and/or amygdalar dys-function might misinterpret environmental cues,such as the facial expressions of others, and reactimpulsively, as a preemptive strike, to a misper-ceived threat. The perception of whether a stim-ulus is threatening is decisive in the cognitiveprocessing leading to the aggressive behavior.Evidence suggests that individuals vary consider-ably in their ability to suppress negative emotion.Therefore, individuals with decreased prefrontalactivity may have greater difficulty suppressingnegative emotions than those individuals whohave greater prefrontal activation. Finally, althoughprefrontal activity helps one to suppress negativeemotion, this negative emotion is generated bysubcortical structures, including the amygdala.Therefore, an individual may be more prone toviolence in general, and impulsive violence, inparticular, if prefrontal functioning is dimin-ished in relation to subcortical activity.

Research on individuals who have sufferedtraumatic head injury is of key importance inunderstanding the neural substrates of aggressiveand/or violent behavior; however, Brower andPrice (2001) noted many limitations of headinjury studies, such as inadequate controls forknown risk factors, including prior history ofaggressive or violent behavior, socioeconomicstatus, stability of employment, and substanceabuse. Research of behavior following headinjury is also one step removed from the questionof whether aggressive and/or violent individuals(who may have no history of head trauma) haveneurological dysfunction localized to specificareas in the brain.

Studies of aggressive, violent, and/or antiso-cial offenders using functional (SPECT and PET)and structural (MRI) neuroimaging are begin-ning to reveal abnormalities in these groups(Raine, Lencz, et al., 2000). Specifically, 17 neu-roimaging studies have been conducted onsamples derived from forensic settings, prisons,

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Section 8 � Neuroimaging Studies of Aggressive and Violent Behavior 301

psychiatric hospitals, and on violent offenderswho are noninstitutionalized. Our review ofthese works reveals four consistent patterns:(a) prefrontal dysfunction is associated withaggressive and/or violent behavioral histories;(b) temporal lobe dysfunction, particularly left-sided medial-temporal (subcortical) activity, isassociated with aggression and/or violence;(c) the relative balance of activity between theprefrontal cortex and the subcortical structures isassociated with impulsive aggression and/or vio-lence; and (d) the neural circuitry underlying theregulation of emotion and its affiliated behaviorsis complex. Each of these patterns is described intheoretical context below.

Prefrontal Dysfunction Is Associated With Aggressive and/or Violent Behavioral Histories

Of the 17 studies reviewed, 14 specificallyexamined possible links between frontal lobepathology and aggressive and/or violent behav-ior. In the 10 SPECT and PET studies, 100%reported deficits in either prefrontal (8 of 10studies) or frontal (2 of 10 studies) functioningin aggressive, violent, and/or antisocial groupscompared to nonaggressive patients or healthycontrols. Analyses of specific regions in themedial prefrontal cortex revealed that individualswho were aggressive and/or violent had signifi-cantly lower prefrontal activity in the OFC (4 of10 studies), anterior medial cortex (5 of 10 stud-ies), medial frontal cortex (2 of 10 studies) and/or superior frontal cortex (1 of 10 studies). In thefour MRI studies, 50% (2 of 4 studies) reporteddecreased grey matter volume in prefrontal orfrontal regions, and 25% (1 of 4 studies) reportednonspecific white matter abnormalities, notlocalized to the frontal cortex.

The consistency with which prefrontal dis-ruption occurs across studies, each of whichinvestigated participants with different types ofviolent behaviors, suggests that prefrontal dys-function may underlie a predisposition to vio-lence. Evidence is strongest for an association

between prefrontal dysfunction and an impulsivesubtype of aggressive behavior. Empirical find-ings concerning the regulation of negative emo-tion suggest that prefrontal sectors, such as theOFC, appear to play a role in the interpretationof environmental stimuli and the potential fordanger. Consequently, disruptions in prefrontalfunctioning may lead individuals who are impul-sive and aggressive to misinterpret situations asthreatening and potentially dangerous, which inturn increases the probability of violent behavioragainst a perceived threat.

Nevertheless, four caveats are noteworthy.First, although prefrontal disruption was consis-tently related to aggressive and/or violent behav-ior, this association may reflect a predispositiononly, requiring other environmental, psychologi-cal, and social factors to enhance or diminish thisbiological risk. Second, prefrontal dysfunctionhas also been documented in a wide variety ofpsychiatric and neurological disorders not asso-ciated with violence, and it may be argued thatfrontal hypometabolism is a general, nonspecificfinding associated with a broad range of condi-tions. However, Drevets and Raichle (1995)reported that although frontal deficits have beenobserved in conditions, such as major depres-sion, schizophrenia, and obsessive-compulsivedisorder, the neurological profile for individualswho are aggressive and/or violent is differentfrom these other groups. For example, whilemurderers exhibit widespread bilateral prefrontaldysfunction, individuals with depression tend tohave disruptions localized in the left hemisphereonly and to the left dorsolateral prefrontal cortex,in particular. (See Raine, Buchsbaum, et al., 1997,for a discussion of alterations in brain function-ing across a variety of psychiatric conditions.)

Of the 10 SPECT and PET studies reviewed,70% reported temporal lobe dysfunction inaggressive and/or violent groups, with reductionsin left temporal lobe activity in 6 of 7 studies.Examination of the medial-temporal lobe, whichincludes subcortical structures, such as the amyg-dala, hippocampus, and basal ganglia, revealedthat subcortical disruptions also characterized

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individuals who were aggressive and/or violent(4 of 7 studies). In the six MRI studies that exam-ined the possibility of temporal lobe abnormali-ties, 100% (6 of 6 studies) reported temporalirregularities, including asymmetrical gyral pat-terns in the temporal-parietal region, decreasesin anterior-inferior temporal lobe volume(including the amygdala-hippocampal region oradjacent areas), increases in left temporal lobevolume, or pathologies specific to the amygdala.

It is important to note that excessive rightsubcortical activity or abnormal temporal lobestructure was most common in patients with ahistory of intense violent behavior, such as thatseen in those with intermittent explosive disor-der rather than in patients who had aggressivepersonality types or who had high scores on anaggression scale. In humans, right-hemisphereactivation has been suggested to play a role in thegeneration of negative affect. Therefore,increased subcortical activity in the right hemi-sphere could lead an individual to experiencenegative affect that promotes aggressive feelingsand acts as a general predisposition to aggressionand violence. These findings are generally consis-tent with current conceptions of emotion regula-tion and its purported relationship to impulsiveviolence, in particular.

The Relative Balance of Activity Between the Prefrontal Cortex and the SubcorticalStructures Is Associated With ImpulsiveAggression and/or Violence

Previous research has suggested that individ-uals may be predisposed to impulsive violence ifprefrontal functioning is diminished relative tosubcortical activity. Raine, Meloy, et al. (1998)found that reduced prefrontal functioning rela-tive to subcortical functioning was characteristicof those who commit impulsive acts of aggres-sion and/or violence. By contrast, aggressionand/or violence of a predatory nature was notrelated to reduced prefrontal and/ or subcorticalratios. They also suggest that although most bio-logical studies of aggression and/or violence have

not distinguished between impulsive and pre-meditated aggression, this distinction is likelyrelevant to understanding the neuroanatomicaland functional underpinnings of these behaviors.

An additional line of evidence that lendssupport to the impulsive and/or predatory dis-tinction comes from investigations regarding themechanism underlying the suppression of nega-tive emotion. The neurochemical link mediatingprefrontal and/or subcortical interactions is pur-portedly an inhibitory serotonergic connectionfrom the prefrontal cortex to the amygdala. Theprefrontal cortex is a region with a high densityof serotonin receptors, which sends efferents tothe brainstem where most of the brain’s serotonin-producing neurons originate. The prefrontal cor-tex, amygdala, and hippocampus also receiveserotonergic innervation. Therefore, it is logicalthat dysfunction in the prefrontal and/or subcor-tical regions disrupts serotonergic activity in thebrain. Consistent with this hypothesis, the sero-tonergic system has been shown to be dysfunc-tional in victims of violent suicide attempts,impulsive violent offenders, impulsive arsonists,violent offender and arson recidivists, childrenand adolescents with disruptive behavior disor-ders, and “acting out” hostility in normal volun-teers. In all those studies, low serotonin levelswere strongly related to the maladaptive behav-iors noted.

y Implications for Criminology and Criminal Justice

Historically, paradigms guiding criminologicalprograms of study have tended to bypass com-plex webs of interconnections that produce andreproduce criminality, favoring instead anemphasis on one dimension or level of analysis.The trend has been to maintain a specializedfocus, often within the confines of a sociologicalor a legalistic model. Attempts to expand theimage of crime through theory integration,which have surfaced quite frequently since themid-1970s, shift attention to different realities ofcrime. The general emphasis, however, has been

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Section 8 � Neuroimaging Studies of Aggressive and Violent Behavior 303

on the integration of ideas within and/or acrossthe two dominant paradigms rather than on abroader, interdisciplinary strategy.

Resistance to interdisciplinarity or discipli-nary cross-fertilization has not been inconsequen-tial. Failure to incorporate interdisciplinaryinsights has stifled exploration of the intersectionsamong structure, culture, and the body, leaving aknowledge void where provocative social facts“merely hang in space as interesting curiosities”(Pallone & Hennessey, 2000, chap. 22, p. 11), andcritical questions go unanswered. More specifi-cally, lack of an imagination of how nurture andnature interact to affect behavior, or what may beunderstood as biography in historical context, hasresulted in an incapacity to either deal with vari-ability or deal with it well. Some male individualssocialized in a patriarchal society rape and someadolescents from poor, urban, single-familieschronically offend; however, most do not. In otherwords, there is individual variation within socialcontexts, and those differences may be betterunderstood if criminologists begin to consider allpertinent angles or dimensions.

Although the studies in our review mayappear to be firmly planted in the tradition ofspecialization and unidimensional thinking, theyshould be interpreted within the framework ofBarak’s (1998) interdisciplinary criminology,where knowledges relevant to a behavioral out-come are treated as complements in an imageexpansion project. Understanding that each per-spective offers a reality of behavior from a differ-ent, though interrelated angle, the objective is todevelop a logical network of theories that willcapture the most dimensions and provide themost accurate information about phenomena ofinterest. In this vein, our appraisal of knowledgefrom the field of neuroscience intends to eluci-date the image of aggression and/or violencewithout supplanting other perspectives and par-adigms. Our desire is not to reduce aggressionand/or violence to brain functioning but toinform of advances in neurological analyses ofemotion regulation and their importance tostudies of that behavior.

It should be noted that the more compre-hensive, interdisciplinary paradigm has beenembraced by some criminologists linked to thebiological sciences. In the 17 studies reviewed,researchers attempted to examine (or at least sta-tistically controlled for) a variety of biological,psychological, and social correlates of aggressiveand/or violent behavior and, in some cases, ana-lyzed biopsychosocial interactions affectingbehavior. Across studies, biological variablesincluded history of head injury, substanceuse/abuse/dependence, diseases of the nervoussystem, left-handedness, body weight, height,head circumference, and sex. Psychological vari-ables included the presence of psychological disorders (such as schizophrenia), indices ofintellectual functioning (such as IQ scores), andperformance-related motivational differences.Social variables included indices of psychosocialdeprivation (such as physical and/or sexualabuse, extreme poverty, neglect, foster homeplacement, being raised in an institution,parental criminality, parental physical fights,severe family conflict, early parental divorce),family size, and ethnicity.

The benefits of integrating ideas or investi-gating an image from several angles in a researchdesign is demonstrated in the neuroimagingstudies provided. Raine, Lencz, et al. (2000), forexample, found that prefrontal and autonomicdeficits contributed substantially to the predic-tion of group membership (antisocial personal-ity disorder vs. control group) over and above 10 demographic and psychosocial measures.The 10 demographic and psychosocial variablesaccounted for 41.3% of the variance. After theaddition of three biological variables into theregression equation (prefrontal gray matter,heart rate, and skin conductance), amount ofvariance explained increased significantly to76.7%, and the prediction of group membershipincreased from 73% to 88.5% classified correctly.These findings suggest that a more contextual-ized theoretical grasp of aggression and/or violenceis possible when this behavior is conceptualizedas multi-dimensional. When nature-nurture

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dichotomies are countered by interdisciplinaryimage expansion, clues about individual variabil-ity emerge, and criminologists come closer tounderstanding the complexity of aggressionand/or violence.

The compelling evidence about this behav-ior revealed in the reviewed neuroimaging stud-ies is valuable, then, not because it allows forcompletely reliable predictions of behavioraloutcomes, but because it makes the image ofaggression and/or violence a little less murky.Moreover, when merged with existing knowl-edges, particularly ideas about social structuresand social psychology (sociological model) andrational choice (legalistic model), such findingsmay spawn new visions of justice centered onprevention and treatment. Within an interdisci-plinary framework that values neuroscience,virtually every essential sociological factor elab-orated by criminologists, structural and proces-sual, acquires a greater potential to explainaggression and/or violence and influence policymaking. According to the works in our review, aswell as other research in this area, all forms ofchild abuse and neglect, direct exposure to vio-lence (including media violence), an unstablefamily life, poor parenting, lack of prenatal andperinatal services, individual drug use, maternaldrug use during pregnancy, poor educationaland employment structures, poverty, and evenexposure to racism play a vital role in the pro-duction of aggression and/or violence. Thus, theinclusion of insights from neuroscience furtherlegitimizes prevention strategies touted by advo-cates of the sociological paradigm, from socialdisorganization theory to self-control theory.

When aggression and/or violence is not pre-vented, the criminal justice system is grantedresponsibility for social control. . . . Drawingfrom empirical findings across disciplines andlevels of analysis, a vision of therapeutic justiceencourages the development of holistic treat-ment regimens that hold offenders to “scientifi-cally rational and legally appropriate degree[s] ofaccountability” (Nygaard, 2000, chap. 23, p. 12).

The potential for this approach to replace theutilitarian model lies in its continued ability tounveil the often-perplexing ways in which choiceis structured. This facilitates an awareness thatthe legally appropriate and the scientificallyrational are in unity. Human creativity is notignored in this paradigm; however, the clearerimage it provides points to an amalgam of limi-tations. When an individual is brought into thecriminal justice system, an inter-disciplinarianseeks to examine those restrictions on behaviorand to tailor treatments accordingly.

With varying levels of success, criminolo-gists have sought to qualify choice and diminishthe impact of legalistic factors on conceptions ofjustice since the advent of positivism in the 19thcentury. Assessments of measures associatedwith social psychology, psychology, and psychia-try, along with input implicating structuralconcerns, such as unemployment, have been uti-lized, and a plethora of interventions haveevolved. Thus, cracks in the utilitarian mold ofjustice have accrued, laying the foundation forinterdisciplinarity in thought and in treatment.Applied to aggression and/or violence, thistranslates into the implementation of treatmentplans with multidimensional components, toinclude neurological techniques that addresshow brain dysfunction affects choice. Althoughnot the sole neurological strategy, the interven-tion most consistently promoted is drug therapy.Several types of drugs, such as anti-convulsants,psychostimulants, and serotonergic agents, havebeen successful in reducing aggressive behavior.Inter-disciplinary thinkers should not be hesi-tant to consider using these pharmacologicalremedies when biopsychosocial indicators over-whelmingly suggest that an individual is at riskto violently recidivate, for it is a step in the direc-tion of therapeutic justice.

Other than paradigmatic preferences disal-lowing an interdisciplinary consideration ofaggressive and/or violent crimes and lack offunding, the largest obstacle in attaining thera-peutic justice is the inability to predict future

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Section 8 � Neuroimaging Studies of Aggressive and Violent Behavior 305

behavior. When informed by neuroscience, clas-sification and prediction instruments are fine-tuned. To illustrate, Robinson and Kelley (2000)discovered that, among probationers, indica-tors of brain dysfunction correlated with repeatviolent offending, as opposed to repeat nonvio-lent offending and first-time offending. Birthcomplications, family abuse, head injury,parental drug use, abnormal interpersonal char-acteristics, and offender substance abuse werefound to be risk factors for recidivism within thisgroup. Given that it is estimated that less com-prehensive prediction models reap false positivesin approximately two thirds of all cases, addedprecision is welcome.

Still, prediction is not foolproof. Short-comings in this area lead some to conclude thatdrug therapy and other invasive strategies areunwarranted. Before throwing in the towel, itshould be acknowledged that pharmacologicalremedies already abound in the criminal justicesystem, along with many other intrusions.Knowledge from neuroscience merely allows forthe targeted distribution of services to appropri-ate populations, a fruitful strategy given thescarcity of resources at the system’s disposal.Prevention strategies directed at alleviating envi-ronmental conditions that increase the probabil-ity for aggression and/or violence are optimal;however, criminologists should not dismiss neu-roscientific individual-level interventions incases where patterns of aggressive and/or violentcriminality are detected. Converging lines of evi-dence suggest that those patterns are producedby a unique combination of external and internalrisk factors, each of which is integral to the con-struction of treatment regimens intended toeffect therapeutic justice.

Blind spots in the image of aggression and/or violence should not deter interventions wherethey hold promise for enhancing quality of life. Itis unfair and unjust to those processed in thecriminal justice system and to society at large forcriminologists to ignore this evidence and thecontrol strategies proposed.

y Conclusion

Functional and morphometric neuroimaginghave enhanced our understanding of the dis-tributed neural networks that subserve com-plex emotional behaviors. Research emanatingfrom affective, behavioral, and clinical neuro-science paradigms is converging on the conclu-sion that there is a significant neurologicalbasis of aggressive and/or violent behavior overand above contributions from the psychosocialenvironment. In particular, and consistentwith modern theories of emotion regulation,reduced prefrontal and/or subcortical ratiosmay predispose to impulsive aggression and/orviolence. Further progress in the study of thesebehaviors will require a forensically informed,interdisciplinary approach that integratesneuropsychological and psychophysiologicalmethods for the study of the brain, emotionalprocessing, and behavior.

As this line of interdisciplinary researchunfolds, it is vital that criminology and criminaljustice begin to incorporate what is knownabout human behavior into its explanatorymodels, as well as its classrooms. Evidence sug-gests that brain structure and brain functioningdo affect behavior, particularly aggressive and/or violent behavior. It is also the case that neuro-science offers means for curbing aggression and/or violence. Traditional criminology and crimi-nal justice paradigms tend to sidestep these issuesbecause of aversions to less dominant knowl-edges, especially biological programs of study.Biological insights are often dubbed Lombrosian,suggesting that some behavioral scientists retainnotions of a born criminal easily identifiableusing some magic test. Continued aversion toanything biological on these grounds is anachro-nistic and will hamper the development oftheory and policy.

The problem is that neurobiological dis-covery has carried on with little to no inputfrom criminology and criminal justice, andthere is every reason to believe that the research

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will progress. There is also reason to believe thatthe functioning of the criminal justice systemwill be affected by the findings produced. Thegeneral public is already being widely exposedto such advances through numerous televisionnews clips and articles appearing in newspapersand weekly periodicals. If criminology andcriminal justice wants to be relevant in morethan a historical sense when it comes to theoriz-ing about aggressive and/or violent behaviorand formulating policies accordingly, it isimperative that the field embrace the interdisci-plinary model.

y Implications for Practice,Policy, and Research

Practice

Bridging the gap between nature and nur-ture, a biopsychosocial model for understandingaggression enhances the explanatory capacity ofsociologically based criminological theories byaccounting for individual variability withinsocial contexts.

Insights derived from a biopsychosocialmodel offer the most promise in the realm ofcrime prevention, which entails devising holistictreatment strategies for those exposed to numer-ous risk factors.

Policy

The accuracy of risk classification devices,used extensively throughout the criminal justice process, may be enhanced by incorporat-ing what is known about negative emotion regulation.

Research

Research reveals that other cortical and sub-cortical structures likely play a role in emotionregulation through their inextricable link to theprefrontal and medial-temporal regions. Thecomplexity of this neural circuitry must beexplored with greater precision.

y References

Barak, G. (1998). Integrating criminologies. Needham Heights,MA: Allyn & Bacon.

Blair, R. J. R., Morris, J. S., Frith, C. D., Perrett, D. I., & Dolan,R. J. (1999). Dissociable neural responses to facial expres-sions of sadness and anger. Brain, 122, 883–893.

Brower, M. C., & Price, B. H. (2001). Neuropsychiatry offrontal lobe dysfunction in violent and criminal behav-ior: A critical review. Journal of Neurology, Neurosurgery,and Psychiatry, 71, 720–726.

Davidson, R. J., Putnam, K. M., & Larson, C. L. (2000).Dysfunction in the neural circuitry of emotion regulation:A possible prelude to violence. Science, 289(5479), 591–594.

Dougherty, D. D., Shin, L. M., Alpert, N. M., Pitman, R. K.,Orr, S. P., Lasko, M., et al. (1999). Anger in healthy men:A PET study using script-driven imagery. BiologicalPsychiatry, 46, 466–472.

Drevets, W. C., & Raichle, M. E. (1995). Positron emissiontomographic imaging studies of human emotional disor-ders. In M. S. Gazzaniga (Ed.), The cognitive neurosciences(pp. 1153–1164). Cambridge, MA: MIT Press.

Nygaard, R. L. (2000). The dawn of therapeutic justice. In D. H.Fishbein (Ed.), The science, treatment, and prevention ofantisocial behaviors: Application to the criminal justice system(chap. 23, pp. 1–18). Kingston, NJ: Civic Research Institute.

Pallone, N. J., & Hennessy, J. J. (2000). Indifferent communi-cation between social science and neuroscience: The caseof “biological brain-proneness’ for criminal aggression.In D. H. Fishbein (Ed.), The science, treatment, and pre-vention of antisocial behaviors: Application to the criminaljustice system (chap. 22, pp. 1–13). Kingston, NJ: CivicResearch Institute.

Pietrini, P., Guazzelli, M., Basso, G., Jaffe, K., & Grafman, J.(2000). Neural correlates of imaginal aggressive behaviorassessed by positron emission tomography in healthysubjects. American Journal of Psychiatry, 157(11), 1772–1781.

Raine, A., Buchsbaum, M., & LaCasse, L. (1997). Brain abnor-malities in murderers indicated by positron emissiontomography. Biological Psychiatry, 42, 495–508.

Raine, A., Lencz, T., Bihrle, S., LaCasse, L., & Colletti, P.(2000). Reduced prefrontal gray matter volume andreduced autonomic activity in antisocial personality dis-order. Archives of General Psychiatry, 57, 119–127.

Raine, A., Meloy, J. R., Bihrle, S., Stoddard, J., LaCasse, L., &Buchsbaum, M. S. (1998). Reduced prefrontal andincreased subcortical brain functioning assessed usingpositron emission tomography in predatory and affectivemurderers. Behavioral Sciences and the Law, 16, 319–332.

Robinson, M., & Kelley, T. (2000). The identification of neu-rological correlates of brain dysfunction in offenders byprobation officers. In D. H. Fishbein (Ed.), The science,treatment, and prevention of antisocial behaviors:Application to the criminal justice system (chap. 12,pp. 12-1–12-20). Kingston, NJ: Civic Research Institute.

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Section 8 � A Theory Explaining Biological Correlates of Criminality 307

D I S C U S S I O N Q U E S T I O N S

1. How might we make the case that the findings presented in this review support Agnew’s “supertraits” theory discussed in Section 9?

2. What are the authors’ main arguments for “biopsychosocial” integration?

3. What do you think are the main obstacles to biopsychosocial integration?

READING

A Theory Explaining Biological Correlates of Criminality

Lee Ellis

In this article, Lee Ellis proposes a biosocial theory he calls the evolutionary neuroandrogenictheory (ENA), which purports to explain many of the correlates of criminal behavior. Twopropositions form the theory’s foundation: an evolutionary proposition and a neurohormonalproposition. According to the first proposition, males have been favored for victimizing othersbecause doing so has helped them to compete and acquire resources that they use to attractmates. The second proposition is concerned with identifying the neurochemistry underlying thisevolved strategy. It maintains that male sex hormones—particularly testosterone—alter brainfunctioning in ways that promote competitive and victimizing types of behavior, which includes,but is not limited to, many types of criminality.

SOURCE: Ellis, L. (2005). A theory explaining biological correlates of criminality. European Journal of Criminology, 2(3), 287–315.Reprinted with permission of Sage Publications, Ltd.

Despite growing evidence that biology plays animportant role in human behavior, most theoriesof criminal behavior continue to focus on learningand social environmental variables. This articleproposes a biosocial theory of criminality thatleads one to expect variables such as age, genderand social status will be associated with offendingin very specific ways. According to the theory,androgens (male sex hormones) have the ability to

affect the brain in ways that increase the probabil-ity of what is termed competitive/victimizing behav-ior (CVB). This behavior is hypothesized to existalong a continuum, with “crude” (criminal) formsat one end and “sophisticated”(commercial) formsat the other. Theoretically, individuals whosebrains receive a great deal of androgen exposurewill be prone toward CVB. However, if they havenormal or high capabilities to learn and plan, they

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will transition rapidly from criminal to non-criminal forms of the behavior following the onsetof puberty. Individuals with high androgen expo-sure and poor learning and planning capabilities,on the other hand, often continue to exhibit crim-inality for decades following the onset of puberty.

y The EvolutionaryNeuroandrogenic Theory of Criminal Behavior

The theory to be presented is called the evolu-tionary neuroandrogenic theory (ENA). The maintypes of offenses it attempts to explain are thosethat harm others, either by injuring them physicallyor by depriving them of their property. Two mainpropositions lie at the heart of ENA theory. Thefirst addresses evolutionary issues by assertingthat the commission of victimful crimes evolvedas an aspect of human reproduction, especiallyamong males. The second is concerned withidentifying the neurochemistry responsible forincreasing the probability of criminality amongmales relative to females. The theory maintainsthat sex hormones alter male brain functioningin ways that promote CVB, which is hypothe-sized to include the commission of violent andproperty crimes.

The concept of CVB is illustrated in Table 8.2.At one end of the continuum are acts that inten-tionally and directly either injure others or dis-possess them of their property. In all societies

with written laws, these obviously harmful actsare criminalized. At the other end of the CVBcontinuum are acts that make no profits on thesale of goods or services, although those whoadminister and maintain the organizationsunder which they operate usually receive muchhigher wages than do those who provide most ofthe day-to-day labor. In a purely socialist econ-omy, the latter type of minimally competitiveactivities is all that is allowed; all other forms arecriminalized. A capitalist economy, on the otherhand, will permit profit-making commerce andoften even tolerate commerce that involves sig-nificant degrees of deception. With the conceptof CVB in mind, the two propositions uponwhich the theory rests can now be described.

y The Evolutionary Proposition

Throughout the world, males engage in victim-ful crimes (especially those involving violence)to a greater extent than do females. To explainwhy, ENA theory maintains that female matingpreferences play a pivotal role. The nature ofthis mating preference is that females considersocial status criteria much more than males doin making mate choices, a pattern that hasbeen documented throughout the world (Ellis,2001). From an evolutionary standpoint, thisfemale preference has served to increase thechances of females mating with males who are

308 INTRODUCTION TO CRIMINOLOGY

The Continuum very crude-------------------------------intermediate----------------------------------------very sophisticated

Probability virtually certain-------------------------intermediate-------------------------------------exceedingly unlikelyof Being Criminalized

Examples Violent and Embezzlement, Deceptive Profit- Nonprofit-property fraud (“white business making making offenses collar crime”) practices, commerce commerce(“street crime”) price gouging

Table 8.2 Continuum of Victimizing Behavior (Reflecting Competitive/Victimizing Tendencies)

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Section 8 � A Theory Explaining Biological Correlates of Criminality 309

reliable provisioners of resources, allowingfemales to focus more of their time and energyon bearing offspring. Another consequence hasbeen that female choice has made it possiblefor males who are status strivers to pass ontheir genes at higher rates than males who arenot. Such female preferences are found inother mammals, as evidenced by their matingmore with dominant males than with subordi-nate males.

According to ENA theory, female prefer-ences for status-striving males have caused mostmales to devote considerable time and energyto competing for resources, an endeavor thatoften victimizes others. In other words, naturalselection pressure on females to prefer status-striving mates has resulted in males with aninclination toward CVB. ENA theory main-tains that the brains of males have beenselected for exhibiting competitive/victimizingbehavior to a greater extent than the brains offemales, and that one of the manifestationsof this evolved sex difference is that males are more prone than females toward victimfulcriminality.

Theoretically, the same natural selectionpressure that has resulted in the evolution ofCVB has also favored males who flaunt and evenexaggerate their resource-procuring capabilities.More unpleasant consequences of the female biasfor resource provisioning mates are male tenden-cies to seek opportunities to circumvent femalecaution in mating by using deceptive and evenforceful copulation tactics. This implies that rapewill always be more prevalent among males thanamong females. ENA theory also leads one toexpect complex social systems to develop in orderto prevent crime victimization. In evolutionaryterms, these systems are known as counter-strategies.An example of a counter-strategy to crude formsof CVB is the evolution of the criminal justicesystem.

As with any theory founded on neo-Darwinian thinking, ENA theory assumes thatgenes are responsible for substantial proportionsof the variation in the traits being investigated. In

the present context, the average male is assumedto have a greater genetic propensity toward CVBthan is true for the average female. However, thisassumption must be compromised with the factthat males and females share nearly all of theirgenes. Consequently, the only possible way forthe theory to be correct is for some of the genesthat promote criminality (along with otherforms of CVB) to be located on the one chromo-some that males and females do not share—theY-chromosome.

y The NeuroandrogenicProposition

The second proposition of ENA theory assertsthat three different aspects of brain functioningaffect an individual’s chances of criminal offend-ing by promoting CVB. Two additional neuro-logical factors help to inhibit offending byspeeding up the acquisition of sophisticatedforms of CVB. Testosterone’s ability to affectbrain functioning in ways that promote CVB isnot simple, but most of the complexities will notbe considered here. The main point to keep inmind is that testosterone production occurs intwo distinct phases: the organizational (or perina-tal) phase and the activational (or postpubertal)phase. Most of the permanent effects of testos-terone effects occur perinatally. If levels oftestosterone are high, the brain will be masculin-ized; if they are low, the brain will remain in itsdefault feminine mode.

ENA theory asserts that androgens increasethe probability of CVB by decreasing an individ-ual’s sensitivity to adverse environmental con-sequences resulting from exhibiting CVB. Thislowered sensitivity is accomplished by incliningthe brain to be suboptimally aroused. Suboptimalarousal manifests itself in terms of individualsseeking elevated levels of sensory stimulationand having diminished sensitivity to pain.

The second way androgens promote CVBaccording to ENA theory is by inclining the limbicsystem to seizure more readily, especially under

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stressful conditions. At the extreme, these seizuresinclude such clinical conditions as epilepsy andTourette’s syndrome. Less extreme manifestationsof limbic seizuring are known as episodic dyscon-trol and limbic psychotic trigger. These latter pat-terns include sudden bursts of rage and othernegative emotions, which often trigger forcefulactions against a perceived provocateur.

Third, ENA theory asserts that androgenexposure causes neocortical functioning to beless concentrated in the left (language-dominated)hemisphere and to shift more toward a righthemispheric focus. As a result of this so-calledrightward shift in neocortical functioning, malesrely less on language-based reasoning, emphasiz-ing instead reasoning which involves spatial andtemporal calculations of risk and reward proba-bilities. Coinciding with this evidence areintriguing new research findings based on func-tional magnetic resonance imaging (fMRI) whichsuggest that empathy-based moral reasoningoccurs primarily in the left hemisphere.Predictably, empathy-based moral reasoningseems to be less pronounced in males than infemales. Such evidence suggests that empathy-based moral reasoning is more likely to preventvictimful criminality than so-called justice-basedmoral reasoning.

Theoretically, the three androgen-enhancedbrain processes just described have evolved inmales more than in females because theseprocesses contribute to CVB. Furthermore, com-petitive/victimizing behavior has evolved inmales more than in females because it facilitatesmale reproductive success more than it facilitatesfemale reproductive success.

y Inhibiting Criminal Forms of Competitive/VictimizingBehavior

Regarding the inhibiting aspects of brain func-tioning, two factors are theoretically involved.One has to do with learning ability and the otherentails foresight and planning ability. According

to ENA theory, the ability to learn will correlatewith the rapidity of male transitioning fromcrude to sophisticated forms of CVB. This meansthat intelligence and other measures of learningability should be inversely associated with persis-tent involvement in criminal behavior. Likewise,neurological underpinnings of intelligence suchas brain size and neural efficiency should alsocorrelate negatively with persistent offending.These predictions apply only to persistent vic-timful offending, with a much weaker link tooccasional delinquency and possibly none withvictimless criminality.

The frontal lobes, especially their prefrontalregions, play a vital role in coordinating com-plex sequences of actions intended to accomplishlong-term goals. These prefrontal regions tend tokeenly monitor the brain’s limbic region, wheremost emotions reside. Then the prefrontalregions devise plans for either maximizing pleas-ant emotions or minimizing unpleasant ones. Inother words, for the brain to integrate experi-ences into well-coordinated and feedback-contingent strategies for reaching long-termgoals, the frontal lobes perform what has come tobe called executive cognitive functioning. Moralreasoning often draws heavily on executive cogni-tive functioning since it often requires anticipat-ing the long-term consequences of ones actions.

Factors that can impact executive cognitivefunctioning include genetics, prenatal complica-tions, and various types of physical and chemicaltrauma throughout life. According to ENAtheory, inefficient executive cognitive function-ing contributes to criminal behavior. Similarconclusions have been put forth in recent yearsby several other researchers.

To summarize, ENA theory asserts that threeaspects of brain functioning promote competi-tive/victimizing behavior, the crudest forms ofwhich are victimful crimes. At least partiallycounterbalancing these androgen-promoted ten-dencies are high intelligence and efficient execu-tive cognitive functioning. These latter twofactors affect the speed with which individuals

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Section 8 � A Theory Explaining Biological Correlates of Criminality 311

quickly learn to express their competitive/victimizing tendencies in sophisticated ratherthan crude ways. Sophisticated expressions areless likely to elicit retaliation by victims, their rel-atives, and the criminal justice system than arecrude ones. Males with low intelligence and/orwith the least efficient executive cognitive func-tioning will therefore exhibit the highest rates ofvictimful criminal behavior.

y Correlates of Criminal Behavior

Twelve biological correlates of crime with specialrelevance to ENA theory (testosterone, meso-morphy, maternal smoking during pregnancy,hypoglycemia, epilepsy, heart rate, skin conduc-tivity, cortisol, serotonin, monoamine oxidase,slow brainwave patterns, and P300 amplitude)are discussed below.

Testosterone ENA theory predicts that correla-tions will be found between testosterone andCVB. However, the nature of these correlationswill not involve a simple one-to-one correspon-dence between an individual’s crime probabilityand the amount of testosterone in his/her brainat any given point in time. Earlier, a distinctionwas made between the organizational and acti-vational effects of testosterone on brain func-tioning, and that the most permanent andirreversible effects of testosterone occur perina-tally. For this reason alone, testosterone levels cir-culating in the blood stream or in salivafollowing puberty may have little direct correla-tion with neurological levels, especially withineach sex. Therefore, one should not expect tofind a strong correlation between blood or salivalevels of testosterone among, say, 20-year-oldmales and the number of offenses they have com-mitted even though testosterone levels in thebrain at various stages in development are quiteinfluential on offending probabilities.

Numerous studies have investigated thepossible relationship between blood levels orsaliva levels of testosterone and involvement in

criminal behavior, and most have found modestpositive correlations (Maras et al. 2003). Additionalevidence of a connection between testosteroneand aggressive forms of criminality involves arecent study of domestic violence, where offend-ing males had higher levels of saliva testosteronethan did males with no history of such violence(Soler et al. 2000).

Overall, it is safe to generalize that circulat-ing testosterone levels exhibit a modest positiveassociation with male offending probabilities,particularly in the case of adult violent offenses.According to ENA theory, males are more violentthan females, not because of cultural expecta-tions or sex role training, but mainly because oftheir brains being exposed to much higher levelsof testosterone than the brains of females.

Mesomorphy Body types exist in three extremeforms. These are sometimes represented with abulging triangle. Most people are located in thecenter of the triangle, exhibiting what is termed abasically balanced body type. At one corner of thetriangle are persons who are extremely muscular,especially in the upper body, called mesomorphs.Ectomorphs occupy a second corner. Individualswith this body type are unusually slender andnon-muscular. In the third corner, one findsendomorphs, individuals who are overweight andhave little muscularity.

Studies have consistently revealed thatoffending probabilities are higher among indi-viduals who exhibit a mesomorphic body typethan either of the two other extreme body types(e.g., Blackson & Tarter 1994). ENA theoryexplains this relationship by noting that testos-terone affects more than the brain; it alsoenhances muscle tissue, especially in the upperpart of the body.

Maternal Smoking During Pregnancy There isconsiderable evidence that maternal smokingmay lead to an elevated probability of offspringbecoming delinquent (e.g., Rasanes et al. 1998).ENA theory assumes that fetal exposure to

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carbon monoxide and other neurotoxins foundin cigarette smoke disrupt brain development inways that adversely affects IQ or executive cogni-tive functioning, thereby making it more difficultfor offspring to maintain their behavior withinprescribed legal boundaries. However, it is possi-ble that genes contributing to nicotine addictionmay also contribute to criminal behavior. In fact,a recent study reported that the link betweenchildhood conduct disorders (a frequent precur-sor to later criminality) and maternal smokingwas mainly the result of mutual genetic influ-ences (Maughan et al. 2004).

Hypoglycemia Glucose, a type of natural sugar, isthe main fuel used by the brain. The productionof glucose is largely regulated by the pancreas inresponse to chemical messages from a portion of the brain called the hypothalamus. When thehypothalamus senses that glucose levels arebecoming too high or too low, it sends chemicalinstructions to the pancreas to either curtail orincrease production of glucose by regulating theamount of insulin released into the blood system.In most people, this feedback regulatory processhelps to maintain brain glucose at remarkablystable levels. For a variety of reasons, somepeople have difficulty stabilizing brain glucoselevels. These people are said to be hypoglycemic.Dramatic fluctuations in brain glucose can causetemporary disturbances in thoughts and moods,with the most common symptoms being confu-sion, difficulty concentrating, and irritability.

Studies have indicated that hypoglycemiais associated with an elevated probability ofcrime, especially of a violent nature (e.g.,Virkkunen 1986). To explain such a connection,ENA theory draws attention to the importanceof maintaining communication between thevarious parts of the brain in order to controlemotionality. In particular, if the frontal lobesreceive distorted signals from the limbic system,bizarre types of behavioral responses sometimesresult, including responses that are violent andantisocial.

Epilepsy Epilepsy is a neurological disordertypified by seizures. These seizures are tanta-mount to “electrical storms’ in the brain. Whilepeople vary in genetic susceptibilities, seizuresare usually induced by environmental factorssuch as physical injuries to the brain, viralinfections, birth trauma, and exposure to vari-ous chemicals.

The main behavioral symptoms of epilepsyare known as convulsions (or fits), although notall epileptics have full-blown convulsive episodes.Mild epileptic episodes may manifest themselvesas little more than a momentary pause in an on-going activity accompanied by a glazed stare.Seizures that have little to no noticeable debilitat-ing effects on coordinated movement are calledsubconvulsive (or subclinical) seizures. Studies ofhuman populations have shown that epilepsyaffects only about one in every 150 to 200persons. In prison populations, however, theprevalence of epilepsy is around one in 50, atleast three times higher than in the general pop-ulation (e.g., Mendez et al. 1993).

ENA theory can explain the links betweenepilepsy and offending by noting that very basicand primitive emotional responses sometimesemanate from the limbic region of the brain.While seizures in motor control centers are mostlikely to receive a diagnosis of epilepsy, seizuresin the limbic region could provoke very basicsurvival instincts.

Resting Heart and Pulse Rates Heart and pulserates rise in response to strenuous exercise along with stressful and frightening experiences.Studies have shown that on average, the restingheart rate and pulse rate of convicted offendersare lower than those of persons in general (e.g.,Mezzacappa et al. 1997:463). ENA theory wouldaccount for these relationships by stipulating thatboth low heart and low pulse rates are physiolog-ical indicators of suboptimal arousal. Sucharousal levels should incline individuals to seekmore intense stimulation and to tolerateunpleasant environmental feedback to a greater

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Section 8 � A Theory Explaining Biological Correlates of Criminality 313

extent than individuals with normal or superop-timal arousal under most circumstances.

Skin Conductivity (Galvanic Skin Response)Sweat contains high concentrations of sodium,which is a good electrical conductor. A devicecalled a Galvanic Skin Response (GSR) meter wasdeveloped nearly a century ago to monitorpalmer sweat. The GSR works by measuringelectrical impulses passing through our bodiesfrom one electrode to another. Thus, by puttingone’s fingers on two unconnected electrodes of aGSR device, one completes an electrical circuitthrough which imperceptible amounts of elec-tricity flows. Temperature obviously affects howmuch people sweat, but so too do emotions. Themore intense one’s emotions become (especiallythose of fear and anger), the more one willsweat, and thus the stronger will be the readingson the GSR meter.

Numerous studies have examined the possi-bility that persons with the greatest propensitiestoward criminal behavior have distinctive skinconductivity patterns. These studies suggest thatoffenders exhibit lower skin conductivity understandard testing conditions than do people ingeneral (e.g., Buikhuisen et al. 1989; Raine et al.1996). As in the case of heart and pulse rates,ENA theory can account for such findings byhypothesizing that low GSR readings especiallyunder stressful testing conditions are anotherindication of suboptimal arousal.

Cortisol So-called stress hormones are secretedmainly by the adrenal glands during times ofanxiety, stress, and fear. The stress hormone thathas been investigated most in connection withcriminality is cortisol. Most of these studies havesuggested that offenders have below normallevels (e.g., Lindman et al. 1997). As with heartrates and skin conductivity, one could anticipatea low cortisol-high criminality relationship byassuming that low cortisol production even inthe face of stress is another indicator of subopti-mal arousal. This would suggest that offenders

are less intimidated by threatening aspects oftheir environments than are persons in general.

Serotonin Serotonin is an important neurotrans-mitter. When serotonin is relatively active in thesynaptic regions connecting adjacent nerve cells,people typically report feeling a sense of content-ment and calm. Several drugs that have beendesigned to treat depression and anxiety disor-ders operate by either prolonging the presence ofserotonin in the synaptic gaps between neuronsor by facilitating the ability of receptor sites onthe dendrites to bond to the serotonin that isavailable. Low serotonin activity has been linkedto crime by numerous studies, especially impul-sive crimes (e.g.,Virkkunen et al. 1996; Matykiewiczet al. 1997). Explaining the link between sero-tonin and criminality from the perspective ofENA theory draws attention to serotonin path-ways connecting the brain’s prefrontal areas withthe emotion-control centers in the limbic system.Serotonin may facilitate the sort of executive cog-nitive functioning required to restrain impulsivebehavior, especially regarding rage and persistentfrustration.

Monoamine Oxidase Monoamine oxidase(MAO) is an enzyme found throughout the body.Within the brain, MAO helps to break down andclear away neurotransmitter molecules (includ-ing serotonin), portions of which often linger inthe synaptic gap after activating adjacent nervecells. Studies indicate that MAO activity isunusually low among offenders (e.g., Alm et al.1996; Klinteberg 1996). ENA draws attention tothe fact that low MAO activity seems to berelated to high levels of testosterone. Further-more, low MAO brain activity may interferewith the brain’s ability to manufacture or utilizeserotonin.

Brain Waves and Low P300 Amplitude Brainwaves are measured using electrodes placed onthe scalp. These electrodes can detect electricalactivity occurring close to the surface of the brain

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fairly clearly. Despite their complexity, brainwaves can be roughly classified in terms of rang-ing from being rapid and regular (alpha brainwaves) to being slow and irregular (delta brainwaves). Most studies based on electroencephalo-graphic (EEG) readings have found that offend-ers have slower brain waves than do persons ingeneral (e.g., Petersen et al. 1982).

Unlike traditional brain wave measurement,modern computerized brain wave detection isable to average responses to dozens of identicalstimuli presented to subjects at random intervals.This reveals a distinctive brain wave pattern or“signature” for each individual. Nearly everyoneexhibits a noticeable spike in electrical voltage,interrupted by a “dip” approximately one-thirdof a second following presentation of test stimuli.This is called the P300 amplitude of an event-related evoked potential. From a cognitive stand-point, the P300 amplitude is thought to reflectneurological events central to attention andmemory.

While research has been equivocal thus farin the case of criminality, several studies havefound a greater dip in P300 responses by indi-viduals diagnosed with antisocial personalitydisorder than is true for general populations(see Costa et al. 2000). ENA theory can accountfor slower EEG patterns among offenders and aP300 decrement among persons with antisocialbehavior by again focusing on suboptimalarousal. From a neurological standpoint, bothslow brain waves and a tendency toward agreater than normal P300 decrement can beconsidered symptomatic of suboptimal arousal.If ENA theory is correct, both of these condi-tions will be found associated with elevatedbrain exposure to testosterone.

y Summary and Conclusions

Unlike social environmental theories, the evolu-tionary neuroandrogenic (ENA) theory canaccount for statistical associations between

biological variables and criminal behavior.Furthermore, ENA theory predicts the universalconcentration of offending among malesbetween the ages of 13 and 30, patterns thatstrictly environmental theories have always haddifficulty explaining. As its name implies, ENAtheory rests on two over-arching assumptions.The first assumption is an extension of Darwin’stheory of evolution by natural selection. It main-tains that males on average exhibit CVB morethan females because females who prefer to matewith such males increase their chances of havingmates who are competent provisioners ofresources. These female biases have evolvedbecause females who have had the assistance ofcompetent provisioners have left more offspringin subsequent generations than other females.No comparable reproductive advantage comes tomales who select mates based on resource pro-curement capabilities.

Some forms of CVB are crude in the sense ofrequiring little learning, nearly all of which areeither assaultive or confiscatory in nature. Otherforms are sophisticated in the sense that theyrequire complex learning and involve much moresubtle types of “victimization.” A major expres-sion of sophisticated competitive/victimizingbehavior involves profitable business venturesand/or the management of large organizations.In most societies, these expressions are toleratedand even encouraged. However, the vast majorityof people in all societies condemn the crudestexpressions of CVB, and, in all literate societies,the criminal justice system has evolved to punishsuch behavior.

The theory’s second assumption is thatgenes on the Y-chromosome have evolved whichcause male brains to exhibit higher rates ofcompetitive/victimizing behavior than femalebrains. These genes operate in part by causingwould-be ovaries to develop instead into testesearly in fetal development. Once differentiated,the testes produce testosterone and other sexhormones, which have three hypothesized effects

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Section 8 � A Theory Explaining Biological Correlates of Criminality 315

upon brain functioning, all of which promoteCVB. The three effects are termed suboptimalarousal, seizuring proneness, and a rightward shiftin neocortical functioning. Furthermore, two neu-rological processes are hypothesized to help indi-viduals shift from crude to sophisticated forms ofcompetitive/victimizing behavior. These arelearning ability (or intelligence) and executivecognitive functioning (or planning ability). Thebetter one’s learning ability or executive func-tioning, the quicker he/she will transition fromcrude to sophisticated forms of the behavior.

To illustrate the theory’s predictive power,the second portion of this article reviews evi-dence regarding several biological correlates ofcriminal behavior. For all of these correlates,the theory was able to explain their apparentrelatedness to criminality. Proponents of criti-cal theory, differential association theory,control theory, etc. cannot explain these corre-lates. They must simply ignore them. Overall,ENA theory should help move criminologybeyond strictly social environmental theoriestoward a new, more variable-rich paradigm.From the theory’s perspective, criminalityresults from a complex interaction of evolu-tionary, biological, learning, and social envi-ronmental factors.

y References

Alm, P. O., af Klinteberg, B., Humble, K., Leppert, J.,Sorensen, S., Thorell, L. H., et al. (1996). Psychopathy,platelet MAO activity and criminality among formerjuvenile delinquents. Acta Psychiatrica Scandinavica, 94,105–111.

Blackson, T. C., & Tarter, R. E. (1994). Individual, family, andpeer affiliation factors predisposing to early-age onset ofalcohol and drug use. Alcoholism: Clinical andExperimental Research, 18, 813–821.

Buikhuisen, W., Eurelings-Bontekoe, E. H. M., & Host, K. B.(1989). Crime and recovery time: Mednick revisited.International Journal of Law and Psychiatry, 12, 29–40.

Costa, L., Bauer, L., Kuperman, S., Porjesz, B., O’Connor, S., &Hesselbrock, V. M. (2000). Frontal P300 decrements,alcohol dependence, and antisocial personality disorder.Biological Psychiatry, 47, 1064–1071.

Ellis, L. (2001). The biosocial female choice theory of socialstratification. Social Biology, 48, 297–319.

Klinteberg, A. (1996). Biology, norms, and personality: Adevelopmental perspective: Psychobiology of sensationseeking. Neuropsychobiology, 34(3), 146–154.

Lindman, R. E., Aromaki, A. S., & Eriksson, C. J. P. (1997).Sober-state cortisol as a predictor of drunken violence.Alcohol and Alcoholism, 32, 621–626.

Maras, A., Laucht, M., Gerdes, D., Wilhelm, C., Lewicka, S.,Haack, D., et al. (2003). Association of testosterone anddihydrotestosterone with externalizing behavior in ado-lescent boys and girls. Psychoneuroendocrinology, 28,932–940.

Matykiewicz, L., La Grange, L., Vance, P., Wang, M., & Reyes, E.(1997). Adjudicated adolescent males: Measures of uri-nary 5-hydroxyindoleacetic acid and reactive hypo-glycemia. Personality and Individual Differences, 22,327–332.

Maughan, B., Taylor, A., Caspi, A., & Moffitt, T. E. (2004).Prenatal smoking and early childhood conduct prob-lems: Testing genetic and environmental explanations ofthe association. Archives of General Psychiatry, 61, 836.

Mendez, M. F., Doss, R. C., & Taylor, J. (1993). Interictal vio-lence in epilepsy: Relationship to behavior and seizurevariables. The Journal of Nervous and Mental Disease, 181,566–569.

Mezzacappa, E., Tremblay, R. E., Kindlon, D., Saul, J. P.,Arseneault, L., Seguin, J., et al. (1997). Anxiety, antisocialbehavior, and heart rate regulation in adolescent males.Journal of Psychiatry, 38, 457–469.

Petersen, K. G. I., Matousek, M., Mednick, S. A., Volovka, J., &Pollock, V. (1982). EEG antecedents of thievery. ActaPsychiatrica Scandinavica, 65, 331–338.

Raine, A., Venables, P. H., & Williams, M. (1996). Better auto-nomic conditioning and faster electrodermal half-recoverytime at age 15 years as possible protective factors againstcrime at age 29 years. Developmental Psychology, 32,624–630.

Rasanes, P., Hakko, H., Isohanni, M., Hodgins, S., Jarvelin,M.-R., & Tiihonen, J. (1999). Maternal smoking duringpregnancy and risk of criminal behavior among adultmale offspring in the Northern Finland 1966 birthcohort. American Journal of Psychiatry, 156, 857–862.

Soler, H., Vinayak, P., & Quadagno, D. (2000). Biosocialaspects of domestic violence. Psychoneuroendocrinology,25, 721–739.

Virkkunen, M. (1986). Reactive hypoglycemic tendencyamong habitually violent offenders. Nutrition Reviews,44(Supplement), 94–103.

Virkkunen, M., Eggert, M., Rawlings, R., & Linnoila, M.(1996). A prospective follow-up study of alcoholic vio-lent offenders and fire setters. Archives of GeneralPsychiatry, 53, 523–529.

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D I S C U S S I O N Q U E S T I O N S

1. All three readings in this section have appealed for the integration of biologically informedand strictly environmental theories of criminology. Given the cascade of evidence supportingthe role of biological factors in criminality, why do you think some criminologists still resistintegration?

2. How would you go about testing ENA theory?

3. What is the primary role of testosterone vis-à-vis criminal behavior?

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