Cesare Lombroso and biological determinism in criminology represent the founding effort to place the study of crime on a scientific basis—identifying measurable physical and biological characteristics that distinguished criminal from non-criminal populations and grounding penal intervention in the nature of the offender rather than the gravity of the offense. This article examines Lombroso’s theoretical framework, its empirical methods, its immediate influence on European and American criminology, and the long trajectory of its legacy in biosocial criminology, eugenics, and the persistent tension between individual-level biological explanation and structural accounts of crime. Situated within the broader History of Criminology, the discussion treats Lombroso neither as a historical embarrassment to be dismissed nor as an anticipator of modern neuroscience to be uncritically celebrated, but as a figure whose work crystallized the positivist criminological project in a specific historical moment and whose influence—direct and indirect—continues to shape debates at the intersection of biology, behavior, and criminal justice.
Introduction
When Cesare Lombroso published L’Uomo Delinquente (Criminal Man) in 1876, he announced a break that would define criminological debate for the next century. The classical tradition—Beccaria, Bentham, the utilitarian reformers—had taken the criminal as given and focused on the rationality of state response. Lombroso’s project was different: to study the criminal himself, to measure and classify his body and mind, and to derive from this empirical investigation a scientific theory of why certain individuals commit crime. Crime, on this account, was not primarily a product of rational calculation, moral failure, or social circumstance; it was a manifestation of a biological condition—the atavism of a criminal type whose physical and psychological constitution placed him at an earlier stage of evolutionary development than the law-abiding majority.
The ambition was scientifically serious by the standards of its time. Lombroso worked as a physician and psychiatrist in Italian prisons and asylums, conducting physical examinations of thousands of inmates, measuring skulls and recording anatomical features, comparing criminal populations with soldiers and the general public. He was applying to criminology the comparative anatomy and evolutionary theory that had transformed biology in the wake of Darwin’s On the Origin of Species (1859) and that were producing new sciences of anthropometry and anthropology across Europe. That his conclusions were empirically wrong in their specifics does not mean his methods were simply foolish; they reflected the state of available scientific tools and the theoretical frameworks that seemed most productive at the moment of their application.
What made Lombroso’s work consequential—and dangerous—was not only its scientific pretensions but the policy implications it carried. If criminals were biologically distinct, then punishment calibrated to the offense made no sense: the appropriate social response was intervention calibrated to the offender’s biological condition, up to and including indefinite incapacitation of the incorrigible. This logic, applied with varying degrees of scientific rigor and moral seriousness across the late nineteenth and early twentieth centuries, contributed to eugenics programs, sterilization laws, and racial theories of crime whose consequences were catastrophic.
Lombroso’s Theoretical Framework
Atavism and the Born Criminal
Lombroso’s central claim was that a significant proportion of criminals—he would eventually estimate roughly one-third—were born criminals: individuals whose biological constitution reflected an atavistic regression to an earlier evolutionary stage of human development. This claim rested on the theory, derived from evolutionary biology and comparative anatomy, that the morphological features associated with earlier human ancestors—heavy brow ridges, large jaws, low cranial capacity, irregular skull formations, extra nipples, prehensile toes—persisted in some modern individuals as inherited throwbacks. The born criminal, physically identifiable by these stigmata of atavism, was constitutionally predisposed to violence, dishonesty, and other behaviors that were normal for primitive humans but criminal in civilized society.
The empirical basis for this claim was Lombroso’s examination of thousands of convicted criminals, beginning with the autopsy of the Calabrian bandit Villella in 1870—an examination that Lombroso later claimed produced the foundational insight of his career when he discovered what he interpreted as an atavistic feature in the bandit’s skull. Subsequent research involved systematic measurement and description of criminal populations in Italian prisons, comparing the frequency of various physical features with those found in military populations and non-criminal civilians. Lombroso found that criminal populations showed higher frequencies of certain morphological features—asymmetries, unusual cranial formations, distinctive ear shapes—and interpreted these differences as evidence of atavistic biological constitution (Lombroso, 1876/2006).
The methodological weaknesses in this research were fundamental, though they were not fully identified until Charles Goring’s systematic refutation in The English Convict (1913). Lombroso’s comparison groups were poorly matched; his measurements were inconsistently recorded; his classification of features as stigmata was often subjective; and he made no attempt to control for the socioeconomic conditions—malnutrition, disease, poverty—that might independently produce the physical features he attributed to atavism. Most fundamentally, he confused correlation with causation: even if criminal populations did show higher frequencies of certain physical features (which Goring’s more careful study disputed), this would not establish that those features caused criminal behavior rather than being jointly caused by some third factor.
Beyond Atavism: Epilepsy, Insanity, and the Criminal Typology
L’Uomo Delinquente underwent five editions between 1876 and 1897, and Lombroso substantially revised and expanded his typology in response to criticism. By the later editions he had developed a more complex taxonomy of criminal types that went beyond the born criminal to include the insane criminal (driven to crime by mental illness), the epileptic criminal (for whom he claimed epilepsy was the biological mechanism underlying atavistic regression), the criminaloid (a larger category of occasional criminals who were pushed into crime by circumstance rather than constitution), and the criminal by passion (whose offenses were driven by intense emotion rather than persistent disposition).
This expanded typology represented a genuine engagement with the criticism that the born criminal category was too narrow to account for the variety of criminal conduct, but it also introduced significant conceptual instability. The criteria for distinguishing born criminals from criminaloids, and both from insane criminals, were never clearly specified, and the typology as a whole was not generated by or tested against independent evidence. It had the character of a classification scheme constructed to accommodate recalcitrant cases rather than a theory with clear predictive implications. Nevertheless, the typological approach—the effort to identify distinct subtypes of offenders defined by their psychological and biological characteristics—influenced forensic psychiatry and criminal classification through much of the twentieth century.
Table 1. Lombroso’s Criminal Typology: Categories, Defining Characteristics, Proposed Etiology, and Policy Implications
| Criminal Type | Defining Characteristics | Proposed Biological Etiology | Proportion of Criminals (Lombroso’s Estimate) | Policy Implication | Subsequent Assessment |
|---|---|---|---|---|---|
| Born Criminal | Physical stigmata of atavism; insensitivity to pain; moral insensibility; recidivism | Atavistic hereditary regression; evolutionary throwback | ~33% of all criminals | Indefinite incapacitation; no point in rehabilitative punishment | Empirically refuted by Goring (1913); no reliable stigmata identified |
| Insane Criminal | Mental illness; absence of criminal intent; post-illness onset | Psychiatric disease rather than atavism | Substantial minority | Asylum rather than prison; medical rather than penal treatment | Contributed to mental health diversion; forensic psychiatry |
| Epileptic Criminal | Epileptic seizures preceding offense; impaired consciousness | Epilepsy as biological mechanism of atavistic regression | Significant overlap with born criminal | Medical treatment alongside incapacitation | Epilepsy-crime link largely discredited; neurological complexity underestimated |
| Criminaloid | No pronounced stigmata; situational offending; less recidivist | Weak constitutional predisposition activated by circumstance | Majority of criminals | Deterrence and situational intervention appropriate | Most consistent with modern interactionist models |
| Criminal by Passion | Intense emotional precipitant; no prior record; remorse | Emotional hypersensitivity; not biological in same sense | Small minority | Mitigating circumstances; reduced sentence | Consistent with modern understanding of heat-of-passion offenses |
Lombroso’s Methods and the Positivist Program
Positivism and the Science of the Criminal
Lombroso’s work was embedded in a broader positivist intellectual program that rejected classical criminology’s abstract philosophical foundations in favor of empirical investigation of criminal behavior and criminal persons. The positivist school—whose other major figures included Enrico Ferri and Raffaele Garofalo—held that crime could be explained scientifically by identifying the biological, psychological, and social factors that produced it, and that criminal justice should be reformed accordingly: replacing punishment calibrated to offense gravity with intervention calibrated to the offender’s degree of danger and amenability to treatment.
Ferri’s contribution was to insist that social and economic factors—Lombroso called them “predisposing causes”—played a larger role in criminal behavior than Lombroso’s biological determinism acknowledged. For Ferri, the born criminal was a limiting case; most criminal conduct resulted from the interaction of individual constitutional factors with social conditions of poverty, inequality, and lack of education. This biosocial formulation—which Ferri called criminal sociology—is structurally continuous with modern biosocial criminology, which similarly argues that genetic and neurobiological factors interact with environmental conditions rather than operating independently of them (Beaver et al., 2014).
Garofalo contributed a jurisprudential dimension: a theory of natural crime, distinguishing offenses that violated universal moral sentiments—what he called probity and piety—from merely conventional legal prohibitions. Natural crimes were those that constituted genuine harm to the moral sense of any civilized community; these alone were appropriate subjects of serious criminal sanction. Garofalo’s distinction anticipated the modern criminological distinction between mala in se and mala prohibita offenses and contributed to debates about what behaviors should be criminalized at all.
The Anthropological Measurement Program
Lombroso’s methodological approach—systematic physical measurement of criminal populations—was part of a broader anthropometric research program that characterized late nineteenth-century science. Alphonse Bertillon’s identification system, which used a standardized series of physical measurements to create a unique identifying profile for known criminals, was adopted by police forces across Europe and America in the 1880s and 1890s. Francis Galton’s development of fingerprint identification drew on similar assumptions about the biological individuality of body characteristics. The measurement of head size and brain volume as indices of intelligence and moral capacity was standard in physical anthropology of the period.
This anthropometric tradition was not simply a mistake awaiting correction; it reflected genuine scientific questions about biological individuality, human variation, and the relationship between physical constitution and behavior that continued to generate research well into the twentieth century. The questions it posed—does heredity shape behavioral predispositions? Do neurological differences correlate with behavioral outcomes?—remain live research questions in biosocial criminology today. What changed was not the questions but the methodological tools available for investigating them: twin studies, molecular genetics, neuroimaging, and longitudinal cohort studies have replaced skull measurement and physical stigmata as the instruments of biological inquiry, and they have produced a more nuanced and empirically defensible account of the relationship between biology and criminal behavior.
Reception and Influence in the United States
American Criminology and the Lombrosian Legacy
Lombroso’s work was received in the United States with a mixture of enthusiasm and critical engagement. American criminologists of the late nineteenth and early twentieth centuries were generally sympathetic to biological approaches to crime, and the emerging discipline of criminal anthropology—led by figures such as Arthur MacDonald, who lobbied unsuccessfully for a federal laboratory of criminal anthropology—took Lombroso’s framework as a productive starting point even when they modified its specific claims. Psychologist Henry Goddard’s studies of the “Kallikak family” (1912) and of feeblemindedness as a cause of crime drew on related assumptions about hereditary transmission of behavioral dispositions, and his work contributed to the broader eugenics movement that produced state-level sterilization legislation in the first decades of the twentieth century.
The sociological tradition that emerged from the University of Chicago in the 1920s represented the most consequential American rejection of Lombrosian biological determinism. Robert Park, Ernest Burgess, Clifford Shaw, and Henry McKay argued that crime was not a product of individual biological defect but of the social ecology of urban neighborhoods—that crime rates varied systematically across space in ways that reflected social disorganization rather than the biological composition of populations. Shaw and McKay’s demonstration that crime rates in specific Chicago neighborhoods remained relatively stable across successive immigrant populations with different national origins was a direct empirical refutation of the racial and biological determinism that had characterized earlier approaches: if crime was a biological characteristic of particular ethnic groups, it should move with those groups as they migrated to better neighborhoods, but the data showed that it did not (Shaw & McKay, 1942).
Goring’s Refutation and Its Limits
Charles Goring’s The English Convict (1913) represented the most systematic empirical attack on Lombroso’s specific claims. Goring studied nearly 3,000 English convicts using rigorous comparative methodology—matching criminal and non-criminal populations on relevant demographic variables—and found no evidence for the physical stigmata of atavism that Lombroso had claimed distinguished criminal populations. His conclusion was unambiguous: “The physical and mental constitution of both criminal and law-abiding persons, of the same age, stature, class, and intelligence, are identical.” There is no such thing as an anthropological criminal type.
Goring’s refutation of the specific Lombrosian claims did not, however, eliminate biological approaches to crime. Goring himself found evidence of hereditary transmission of a general factor of criminal diathesis that he identified with defective intelligence, and he argued that selective reproduction policies—eugenic measures—were the appropriate response. This maneuver—abandoning Lombroso’s morphological determinism while retaining a hereditary framework—was characteristic of the subsequent trajectory of biological criminology: the specific claims of the born criminal theory were abandoned, but the general orientation toward hereditary and constitutional explanations of criminal behavior persisted through new empirical vehicles.
The Lombrosian Legacy: From Eugenics to Biosocial Criminology
Eugenics and the Criminal
The connection between Lombrosian biological determinism and the American eugenics movement was not logically necessary but was historically close. If criminal behavior was substantially hereditary—a claim that biological criminology in various forms consistently advanced—then preventing the reproduction of criminal individuals and families was a rational public health response, on utilitarian grounds. This logic animated the eugenics movement that achieved legislative expression in the United States through sterilization statutes enacted in over thirty states between 1907 and the 1930s, with Virginia’s statute upheld by the Supreme Court in Buck v. Bell (1927). The criminal justice system was explicitly among the target populations for eugenic intervention: the Indiana sterilization law of 1907, the first in the nation, applied to confirmed criminals, idiots, imbeciles, and rapists in state institutions.
The catastrophe of Nazi eugenics—which drew explicitly on American sterilization laws as a model—delegitimized eugenic approaches to crime and effectively ended their open expression in American criminology for several decades. The biological criminology that emerged in the postwar period was more cautious in its claims, more sophisticated in its methodology, and more careful to distinguish empirical findings about statistical associations between biological variables and criminal behavior from normative conclusions about appropriate policy responses. Whether this caution was sufficient to prevent the recapitulation of earlier errors—the racial coding of biological risk factors, the conflation of statistical correlation with individual determinism—remains a live question in contemporary biosocial criminology (Burt & Simons, 2014).
Modern Biosocial Criminology and Lombroso’s Shadow
Contemporary biosocial criminology—which uses twin studies, adoption studies, molecular genetic analysis, and neuroimaging to investigate the biological contributions to criminal behavior—has established empirical findings that are significantly more defensible than Lombroso’s but that raise some structurally similar issues. Twin studies consistently find heritability estimates for antisocial behavior in the range of 40–60%, indicating that genetic factors account for a substantial portion of the variance in criminal and delinquent behavior (Rhee & Waldman, 2002). Neuroimaging studies have found structural and functional differences in prefrontal cortex and amygdala between individuals with persistent antisocial behavior and controls. Gene-environment interaction studies have identified specific genetic variants that moderate the relationship between early adversity and subsequent antisocial behavior.
These findings differ from Lombroso’s in being generated by methods with far greater internal validity, in acknowledging the complexity of gene-environment interaction rather than asserting simple biological determination, and in recognizing that heritability estimates are population statistics that say nothing about the fate of specific individuals. They nevertheless remain subject to the criticisms that have been leveled at biological criminology since Lombroso: that they tend to locate the explanation of crime in characteristics of individuals rather than social structures, that they are vulnerable to racial coding given the racial structure of the populations studied, and that the policy implications they carry must be carefully examined for consistency with individual rights and equal treatment under law (Burt & Simons, 2014).
Lombroso on Gender and the Female Offender
La Donna Delinquente and Its Gender Logic
Lombroso’s theorization of the criminal extended to gender in ways that reveal the ideological assumptions embedded in his ostensibly scientific framework. La Donna Delinquente (Criminal Woman, the Prostitute, and the Normal Woman), published with Guglielmo Ferrero in 1893, applied the atavism framework to female offending with results that were, if anything, more theoretically strained than the born criminal theory itself. Lombroso argued that women were evolutionarily less differentiated than men—biologically less developed, more primitive—and that this greater primitiveness meant that female criminals were less atavistic in the sense relevant to male crime: since all women were closer to the primitive baseline, the female criminal represented a different and in some ways less pronounced form of deviance than her male counterpart.
The logic of this argument produced a characteristically contradictory conclusion: female criminals were simultaneously less atavistic than male criminals and more morally reprehensible, because their deviation from the feminine norm of passivity and nurture represented a more fundamental disruption of natural order. The prostitute—whom Lombroso treated as the female equivalent of the male born criminal—was defined by her sexuality rather than her violence, reflecting the gender-specific content of the deviance categories applied to women. This sexualized construction of female deviance, in which women’s criminal character was understood primarily through the lens of their reproductive and sexual roles rather than through analysis of their actual offending behavior, established patterns of analysis that feminist criminologists would identify and critique a century later.
The practical consequences of the gendered dimension of Lombrosian criminology were significant. Female offenders were understood as sexually deviant rather than dangerous in the way male criminals were, directing institutional responses toward the correction of sexual and moral irregularity rather than the incapacitation or rehabilitation of criminal behavior as such. Reform schools and correctional facilities for women in the late nineteenth and early twentieth centuries reflected this logic: they emphasized domestic training, sexual surveillance, and moral instruction rather than vocational preparation or cognitive rehabilitation. The classification of women’s criminality as primarily a problem of sexual irregularity, rather than a problem of antisocial behavior with social structural roots, obscured both the structural determinants of female offending—poverty, victimization, economic marginalization—and the specific character of the offenses women actually committed.
The Feminist Critique and Its Criminological Legacy
The feminist criminology that emerged in the 1970s mounted a systematic challenge to the gender assumptions embedded not only in Lombrosian biological determinism but in the entire positivist criminological tradition. Freda Adler’s Sisters in Crime (1975) and Rita Simon’s Women and Crime (1975) initiated a research program examining female offending on its own terms rather than as a deviation from male norms. Carol Smart’s Women, Crime and Criminology (1976) provided a more radical theoretical critique, arguing that criminology’s core concepts—including the definition of crime, the construction of the criminal subject, and the theory of causal explanation—were gender-coded in ways that made the discipline incapable of adequately theorizing women’s experience of crime and criminal justice without fundamental reconceptualization.
The feminist critique of Lombrosian criminology specifically identified three recurring problems: the pathologization of female deviance through medicalization and sexualization; the invisibility of women’s victimization in frameworks focused on offending; and the application of gender-specific double standards in criminal justice processing that punished women for sexual and moral irregularity as much as for offense seriousness. These patterns—whose roots feminist criminologists traced to Lombrosian and related biological frameworks—remain empirically documented features of American criminal justice processing that continue to generate research and policy attention (Chesney-Lind & Pasko, 2013).
Lombroso’s Comparative Reception Across National Contexts
Differential National Receptions
Lombroso’s theory was received differently across national criminological traditions in ways that reflected each country’s specific scientific and political culture. In Italy, the positivist school commanded substantial authority through the early twentieth century and influenced both legal doctrine—Italian criminal procedure incorporated expert psychiatric assessment of the accused’s dangerousness—and penal practice. In France, the reception was more critical: Gabriel Tarde’s competing theory of crime as social learning through imitation rather than biological constitution attracted significant following, and the French statistical tradition associated with Lacassagne emphasized social conditions over individual biology.
In Britain, Goring’s systematic empirical refutation of Lombrosian stigmata effectively ended that specific tradition’s influence, though biological approaches persisted in modified forms through the eugenics movement. In the United States, as discussed above, the initial enthusiasm for biological criminology gave way to the sociological approach of the Chicago School, but biological explanations retained institutional presence in the prison classification systems and eugenic programs that expanded through the 1930s. In Germany, Lombrosian biological criminology was eventually synthesized with racial theory in ways that contributed to the catastrophic outcomes described in the article on eugenics and sterilization.
These differential national receptions illustrate that the influence of a scientific theory on policy and practice is not determined by its empirical merits alone but by the specific institutional, political, and cultural contexts in which it is received. The sociology of scientific knowledge—the study of how scientific claims are produced, evaluated, and deployed in specific social contexts—provides an important supplement to the internal history of criminological ideas in explaining why some theories achieve institutional authority and others do not, regardless of their relative empirical support.
The Measurement Debate: From Skulls to Neuroimaging
Instrumentation and the Shifting Targets of Biological Criminology
A recurring feature of biological criminology from Lombroso to the present is that its specific empirical claims have been consistently refuted by improved measurement, while the general orientation toward biological explanation has survived by migrating to new and supposedly more reliable instruments. Lombroso’s skull measurements were refuted by Goring’s more careful anthropometry; Goring’s intelligence deficit claims were undermined by the development of better IQ tests and more rigorous comparative methodology; the chromosomal theories of the 1960s (linking extra Y chromosomes to violence) were discredited by epidemiological research; and EEG studies linking brain wave abnormalities to criminal behavior gave way to the neuroimaging era with its own promises of reliable biological identification of criminal predisposition.
Each instrument generation has repeated a characteristic pattern: initially enthusiastic claims of biological markers distinguishing criminal from non-criminal populations, followed by methodological critique identifying confounds and selection biases in the research designs, followed by more modest and qualified claims that acknowledged the complexity of gene-environment interaction and the impossibility of identifying simple biological determinants of complex behavioral outcomes. Contemporary neuroimaging research on persistent antisocial behavior is more methodologically sophisticated than Lombroso’s skull measurements, but it faces structurally similar challenges: the neural differences associated with antisocial behavior overlap substantially with the neural effects of environmental adversity, trauma, and developmental disadvantage, making it extremely difficult to identify neural predictors of antisocial behavior that are independent of the social conditions that also predict it.
Adrian Raine’s neurocriminology—the most systematic contemporary attempt to develop a biological criminology grounded in neuroimaging evidence—acknowledges these challenges while arguing that the evidence for biological contributions to serious violence is now sufficiently strong to warrant both scientific recognition and cautious policy engagement. Raine’s meta-analyses find consistent structural and functional differences in prefrontal cortex, amygdala, and related structures between individuals with violent antisocial behavior and controls, and he argues that these differences, whether caused by heredity, prenatal adversity, or childhood trauma, constitute meaningful biological risk factors that should inform both prevention and intervention (Raine, 2013). Whether these biological risk factors can be addressed through policy without reproducing the targeting and control implications of earlier biological criminology is the central challenge that neurocriminology must address to avoid Lombroso’s legacy.
The policy implications that Raine draws from neurocriminological evidence are carefully qualified but nonetheless controversial. He advocates early identification of neurological risk factors in childhood, followed by targeted intervention to address modifiable aspects of those risk profiles—nutritional supplementation, early childhood programs designed to improve executive function, and environmental interventions to reduce prenatal and early childhood exposure to neurotoxins that impair brain development. Whether these interventions would in practice be applied with sufficient nuance to avoid the targeting and stigmatization of minority populations—who are disproportionately exposed to the environmental neurotoxins and early adversity that produce neurological risk factors—is a question Raine acknowledges but cannot resolve from within the neurocriminological framework alone.
The history of Lombrosian criminology suggests why this caution is warranted. Every generation of biological criminology has presented its instruments and methods as an improvement over its predecessors, and every generation has been correct in this assessment while simultaneously reproducing structural features—the identification of heritable risk in populations already subjected to structural disadvantage; the conflation of statistical association with individual prediction; the policy application of probabilistic research findings to individual determinations of risk—that characterized the tradition’s most problematic episodes. Contemporary neurocriminology is more sophisticated, more self-aware, and more empirically grounded than Lombroso’s anthropometry; whether that sophistication is sufficient to prevent the recapitulation of Lombroso’s errors in new form is the central question that the tradition’s future history will answer.
The reference list Lombroso’s neurocriminological successors cite is vastly more technically sophisticated than anything he produced, and the findings they report are more reliable. But the fundamental questions he posed—what makes some individuals persistently antisocial while others in similar circumstances are not, what biological processes underlie this difference, and how society should respond to individuals whose biology places them at elevated risk—remain as contested and consequential as they were when he first articulated them in the Italian prisons of the 1860s. That continuity between founding question and contemporary research program is, whatever its empirical vicissitudes, Lombroso’s most significant contribution to the discipline he helped to create.
Conclusion
Cesare Lombroso’s place in criminological history is secure but ambivalent. He founded the positivist criminological program that made the scientific study of criminal behavior—as opposed to philosophical analysis of criminal law—the central project of the discipline. He was wrong about virtually every specific empirical claim he made. His work contributed, through a complex and often indirect chain of influence, to eugenic policies whose consequences were catastrophic. And his fundamental questions—about the biological contributors to criminal behavior, about the relationship between individual constitution and situational circumstance, about the appropriate basis for criminal intervention—remain at the center of criminological debate today.
Contemporary criminology has largely moved away from the individual-level biological determinism that characterized Lombroso’s framework, toward interactionist models that treat biological, psychological, and social factors as jointly shaping the probability of criminal behavior across the life course. But the move toward biosocial explanation—cautious, methodologically sophisticated, empirically constrained—is not a departure from Lombroso’s project so much as its continuation under better conditions. What criminology owes Lombroso is not his specific findings but his insistence that the systematic scientific study of criminal persons is a legitimate and necessary complement to the philosophical analysis of criminal law—an insistence that, once established, proved impossible to abandon even when the specific biological theories it generated were ultimately refuted.
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