Available online at www.sciencedirect.

com

ScienceDirect

The question why and how people differ in personality

cannot be answered satisfactorily while neglecting
biological approaches
Martin Reuter1, Thomas Plieger1 and Petra Netter2

The present opinion paper makes a strong case for a continuum between psychopathology and personality, as
biologically oriented personality theory. A short review on postulated by several authors [2–4]. This will prospec-
seminal studies applying methods from the neurosciences to tively help to improve individual patient centered phar-
unravel the biological underpinnings of personality is given, macological treatment of psychopathologic diseases.
and findings are critically discussed. It becomes apparent
that new methodological advances in the neurosciences
provide inspiration for modern personality research but that Genetic approaches
there is still so much unexplained variance left. Only The nature-nurture debate has addressed exactly the
interdisciplinary research can be a promising research question for the sources of individual differences, and
avenue for the future. quantitative behavioral geneticists decompose observed
Addresses variance in personality traits into genetic and environ-
1
University of Bonn, Department of Psychology, Laboratory of Neuro- mental components [5]. But quantitative genetic research
genetics, Germany (calculating heritability scores by twin studies) is only
2
University of Giessen, Department of Psychology, Germany
statistical by nature, it is not able to provide information
on the biological mechanisms underlying these calculated
Corresponding author: Reuter, Martin ([email protected]) heritability scores. Some personality theories differenti-
ate between temperaments and characters [6]. The for-
mer should be determined to a greater extent by genetic
Current Opinion in Behavioral Sciences 2022, 43:181–186
factors, the latter to a greater extent by environmental
This review comes from a themed issue on Neurobiology of tem-
factors. However, quantitative genetic studies question
perament, personality and psychopathology: what’s next?
this assumption. Additive genetic effects alone provided
Edited by Irina Trofimova and Alan Pickering the most parsimonious explanation for the source of
For complete overview of the section, please refer to the article col- familial aggregation in each character dimension of the
lection, “Neurobiology of temperament, personality and psycho- temperament and character inventory (TCI) [7]. This
pathology: what’s next?”
means that also character is under strong genetic influ-
Available online 17th November 2021 ence. In a similar way, researchers whose theories are
https://doi.org/10.1016/j.cobeha.2021.10.006 based on temperament concepts by Pavlov or Rusalov
2352-1546/ã 2021 Elsevier Ltd. All rights reserved. [8,9] postulate a differentiation between personality and
temperament. Again, only the latter should have a strong
biological basis, as opposed to personality which includes
cultural and social aspects. In our opinion such a rigid
differentiation does not hold empirical testing, since also
personality traits derived by factor analysis like extraver-
Opinion sion or conscientiousness have been shown to comprise
A-biological personality psychology is striving to measure large parts of genetic variance [10–14].
individual differences in personality and temperament by
developing and validating new assessment tools. Most of Not until the late 1990s molecular genetic research on
these approaches use self-report questionnaires, far less of personality started. The invention of the polymerase
these invent paradigms for implicit personality measure- chain reaction (PCR) technique has made this possible.
ment or use experimental approaches [1]. The merits of Since then tremendous technological progress has been
these endeavors are out of question; however, they will made in the analysis of genetic data, starting with single
never be capable to explain underlying biological mech- nucleotide polymorphism (SNP) analysis and culminat-
anisms for the observed differences in personality. That ing in genome wide association studies (GWAS) [15].
is, what drives people to understand why they are how
they are already, the ancient Greek physician Hippo- Genetic data themselves are related to functional and
crates had the idea that differences in temperament have structural differences in brain metabolism and structure
a biological source, the respective dominance of body of the central nervous system. Relating neurotransmitter
fluids. Understanding the biology of personality is not an functionality to individual differences in personality has a
end in itself but it bridges the gap and demonstrates the long tradition. Since prominent personality theories like

www.sciencedirect.com Current Opinion in Behavioral Sciences 2022, 43:181–186

182 Neurobiology of temperament, personality and psychopathology: what’s next?

for example Zuckerman’s Sensation Seeking or Cloninger’s psychoticism, extraversion and neuroticism) make
Bio-Social Personality Theory explain individual differ- straightforward assumptions on the biological basis of
ences in character and temperament by differences in personality. According to Eysenck, high extraversion
monoamines like noradrenaline, dopamine and serotonin should be related to reduced cortical arousal [28]. Such
(see below), molecular genetics focused on candidate an assumption can easily be verified by EEG. For exam-
genes showing associations between these neurotransmit- ple, spectral power derived from resting state EEG was
ter related genes and personality dimensions [16,17]. A used to predict personality traits [29]. The research
broad phenotype like personality is influenced by many question was, if spontaneous oscillations of the brain at
genes, and SNPs can only account for small proportions of rest are characteristic for distinctive personality traits.
variance (often <1%). This is the reason why even meta- This endeavour was successful with respect to the Big
analyses yield conflicting results [18]. On the contrary, Five trait of agreeableness. EEG power ranging from
genome wide association studies (GWAS) are completely 819 Hz was associated with agreeableness across multi-
exploratory by nature and try to relate millions of SNPs to ple recording periods. Results for the remaining Big Five
phenotypic variance. In this endeavor, Lo et al. [19

] traits were less consistent.
conducted a GWAS and identified six SNPS robustly
associated to the Big-Five dimensions. Interestingly, A comprehensive review on the literature testing
these SNPs have also been related to other phenotypes Eysenck’s arousal theory on extraversion summarized
in previous GWAS. For example, Lo et al. found a variant conflicting results and concludes that these heterogenous
associated with neuroticism in L3MBTL2, a gene reported findings are probably due to differences in experimental
to be associated with schizophrenia. These findings again protocols, sample size, or age of subjects. Another possi-
support the continuum model of personality (see below). ble reason is that the studies mainly focused on EEG
power spectral analysis [30]. However, not only resting
Biochemical approaches state EEG but also event-related potentials (ERPs) were
Biochemical explanations for relating neurotransmitter used to test the arousal theory. An innovative approach
activity to personality are rooted in experimental animal was applied in a recent EEG-study using ‘scalp-recorded
research as well as in results from drug treatment in Reward Sensitivity’, a putative marker of dopaminergic
psychopathology. They are based on the concept of a signaling of reward-prediction-error [31
]. The reward-
continuum between temperament and respective psycho- prediction-error hypothesis postulates that midbrain
pathological diseases (e.g. depressive mood ! major dopaminergic neurons signal the discrepancy between
depression, or impulsivity ! impulse control disorders a person’s estimate of the value of an event and the actual
like addiction) [20

]. magnitude of the value. Scalp-recorded Reward Sensitiv-
ity was calculated by contrasting the Feedback-Related-
Unfortunately, brain neurotransmitter activities are not Negativity in response to unpredicted rewards and unpre-
validly represented by their plasma levels. Since, how- dicted non-rewards in an associative learning task. This
ever, all transmitters influence peripheral hormones EEG-marker was clearly related to extraversion and
which can be assessed in blood or saliva, transmitter unrelated to other Big Five personality dimensions prov-
activities can be indirectly inferred from hormone ing discriminant validity.
responses to transmitter related drug challenges
[21,22]. For example, dopamine agonists reliably suppress Imaging approaches
and antagonists increase prolactin release, and respective An example for detecting biological predictors of the
plasma levels may indicate personality related differences propensity to develop a neurological disease by personal-
in dopamine activity [23]. ity traits is given by a neuroimaging (positron emission
tomography (PET)) study investigating the well-known
Each transmitter is involved in several psychological biomarkers for Alzheimer’s disease (AD), the accumula-
functions (emotional, motivational, cognitive), but only tion of tau-proteins and ß-amyloid in the central nervous
a few of these functions, confirmed by different method- system in healthy persons. The study revealed that
ological approaches, are regarded as well-established increased neuroticism was related to higher tau patho-
facts. One of these is that dopamine is involved in reward physiology in cognitively normal participants [32
]. Find-
related functions and traits representing positive emo- ings like this, of course, are primarily of correlational
tionality like extraversion and novelty seeking [24,25]. nature. Two alternative causal explanations for this find-
Furthermore, a consistent finding is the relationship ing are conceivable: either neuroticism predisposes for
between low brain serotonin levels and aggression or lack tau accumulation in the brain or neuroticism changes as a
of impulse control [26,27]. consequence of AD onset, that is, increasing tau levels
may affect personality. Only longitudinal studies can
EEG-studies answer this question of cause and effect, and, finally,
Biologically oriented theories like Eysenck’s PEN-model overlapping sets of genes may be responsible for both tau
(suggesting three factors of personality, namely accumulation and the trait of neuroticism.

Current Opinion in Behavioral Sciences 2022, 43:181–186 www.sciencedirect.com

 The impact of biological personality research Reuter, Plieger and Netter 183

Structural imaging approaches local brain structure and personality are, in part, under
The theoretical assumption that participants scoring high genetic control [40

]. Interestingly, reported findings
on personality dimensions of positive emotionality show could be replicated in two independent samples.
structural differences in the brain’s reward system was
tested and indeed, a positive correlation was observed
between extraversion and gray matter volume in the Genetic imaging
caudate region, a pivotal brain region related to reword It is an acknowledged fact that in most cases prediction
sensitivity [33
]. This voxel-based morphometry (VBM) models are the better the more they rely on multiple
study was based on a large sample of 382 participants of instead of single predictors. Undoubtedly, the same is
whom N = 133 were retested two years later. By means of true for personality. Genetic imaging is such an endeavor
a conjunction analysis it was shown that the body of the to combine two neuroscientific approaches for the study
left caudate nucleus was consistently related to extraver- of temperament. A pioneer study in this respect related
sion scores. The association reported between extraver- the monoamine oxidase A (MAO-A) gene encoding a key
sion and reward related brain morphometry supports enzyme for monoamine catabolism to personality and
theoretical assumptions and empirical work from animal brain activity [41]. Findings showed that carriers of a
research [34]. A study by our own group found a positive low expressing MAO-A allele showed dysregulated amyg-
correlation between extraversion scores and the ratio of dala activation and increased functional coupling with
gray to white matter volume in the left hemisphere [35]. ventromedial prefrontal cortex (vmPFC). Stronger cou-
This association was only detectable in males but not in pling was associated with increased harm avoidance and
females. A speculative explanation for this finding is that decreased reward dependence scores, suggesting that this
language production and processing is located in the left circuitry mediates part of the association of MAO-A with
hemisphere of most people and that communication is a these traits. Interestingly, this finding was again only
key feature of extraversion. Women however, have less observable in males. Which is probably due to the fact
laterality of the brain which is a possible explanation for that the MAO-A gene is located on the X-chromosome,
the null findings in female participants [36]. Another and therefore, in males, there are only monocygotes for
VBM study supported the finding of sex specific associa- this low expressing MAO-A allele. A schematic overview
tions between the Big Five personality traits and gray on how integrative personality research including biolog-
matter volume [37]. Significant associations with gray ical markers works is presented in Figure 1.
matter volume were observable for neuroticism, extraver-
sion, and conscientiousness, however only in male parti- Conclusions
cipants. The authors also speculate that the gender The present mini-review on studies investigating the
effects might be attributable to hormonal interplays or biological basis of individual differences in personality
differences in brain organization between males and demonstrates that new technological advances in the
females. neurosciences provided deeper insights into brain struc-
ture, circuitry and metabolism as related to personality.
Unfortunately, the promising findings by Li et al. [33
]
and others could not be replicated in an even larger It became clear that findings from different disciplines
sample from Israel [38]. In a sample of more than 1100 par- (genetics, neurochemistry, functional and structural neu-
ticipants no associations between the Big Five personality roimaging) differently contributed to elucidating the bio-
traits and variability in brain gray or white matter could be logical basis of specific traits of temperament or person-
found. However, it has to be mentioned that the latter ality. Naı̈ve views that it is possible to map a brain region
study was based on university students limiting the or a neurotransmitter to a certain trait in a 1-to-1 fashion is
generalizability of the findings. obsolete. The brain is organized in complex networks
using identical brain regions, neurotransmitters and genes
Besides the classic voxel-based morphometry analyses, for different emotions and traits. As emphasized by
diffusion tensor imaging (DTI) is a complementary mag- several theories, only the interplay between different
netic resonance imaging (MRI) method providing infor- elements constitutes a specific psychological trait (e.g.
mation on the brain’s networks, for example, the nerve the interplay between different neurotransmitters, neu-
fibers projections. A pioneer work from our group dem- ropeptides and hormones; [28]), or the inspection of
onstrated that individual differences in anxiety are related neuronal networks instead of localized neuronal activity
to white matter tract integrity in the left temporal lobe (e. [42], or the simultaneous analysis of millions of SNPS by
g. a cingulum-hippocampus projection). Again, this find- the technique of GWAS in molecular genetics [19

]. A
ing indicating laterality of the brain was limited to male few studies already combine different methods as shown
participants and could not be found in women [39]. by the approach of genetic imaging. However, this
approach of matching results from different techniques
A recent twin study using data of the Human Connec- and disciplines obtained within the same individuals is
tome Project emphasized that the associations between still rare, but to our opinion, represents the most

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184 Neurobiology of temperament, personality and psychopathology: what’s next?

Figure 1

Genes Environement

Biological Markers
EEG
Personality MRI
Hormones
etc.

Behavior

Surveys Experiments Field Studies

Current Opinion in Behavioral Sciences

An integrative model of biologically influenced personality research. Genes and environment separately and in interaction have an influence on the
brain that controls our behavior moderated by personality. Behavior does not only include overt actions but also motivation, emotions and
cognitions that can be inferred by different methods and by biomarkers.

promising way of better understanding and validating the [43]. However, as Allen et al. posit, ‘personality neurosci-
biological foundation of psychological traits. ence is well-positioned to guide the transition from
description to explanation’ within personality research.
Some personality researchers justify the fact that no
bigger breakthrough in biological personality research The precision in the neuro-methods has been dramati-
has been achieved so far with the missing distinction cally increased — relating the obtained bio-markers to
between socio-cultural influenced character dimensions broad personality traits is admittedly difficult. It is not
and more biologically based temperaments. Especially surprising that the same biomarkers are associated with
the Big Five approach is criticized because of being different phenotypes. In the same way as the complex
developed based on a lexical approach without any theo- configuration of traits constitutes the unique personality
retical assumptions on its biological foundations. We of a distinct individual in the same way interact brain-
agree that personality theories like Gray’s Reinforcement circuits, transmitters and hormones in a complex interplay
Sensitivity Theory (RST) or Cloninger’s neurotransmit- to build up the basis of personality. Read et al. [44]
ter associated Biosocial Theory of Personality are better propose a neural network of the structure and dynamics
starting points for biologically based personality research. of personality integrating the lexical approach, the neuro-
However, one has to admit that the Big-Five’s psycho- biological basis of personality, temperament, goal-based
metric properties, including factor structure and reliabil- models of personality, and an evolutionary analysis of
ities are the best. The factor structure could be replicated motives. This is a critical endeavor to overcome old
in many cultures all over the world and therefore must controversies in personality research adhering either
reflect evolutionary/biological underpinnings. the biological or the lexical approach and the sharp
differentiation between character and temperament.
Obviously, there is a large fraction of traditional person-
ality psychologists who question the merits and perspec- Despite breathtaking knowledge, the question why peo-
tives of a biological oriented personality psychology. ple differ in character and temperament is far from
Their criticism states that neuroscientific approaches only understood. Important information like actual neuro-
helped to describe personality at a lower level of analysis transmitter concentrations can only indirectly be inferred

Current Opinion in Behavioral Sciences 2022, 43:181–186 www.sciencedirect.com

 The impact of biological personality research Reuter, Plieger and Netter 185

by pharmacological challenge tests using endocrinological 7. Gillespie NA, Cloninger CR, Heath AC, Martin NG: The genetic
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Conflict of interest statement of personality: from monoamines to neuropeptides. Neurosci
Nothing declared. Biobehav Rev 2014, 43:228-239 http://dx.doi.org/10.1016/j.
neubiorev.2014.04.006.
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CRediT authorship contribution statement the association between the serotonir transporter gene
polymorphism (5-HTTLPR) and trait anxiety. Mol Psychiatry
Martin Reuter: Conceptualization, Writing – original 2004, 9.
draft. Thomas Plieger: Writing – original draft, Writing
19. Lo M-T, Hinds DA, Tung JY, Franz C, Fan C-C, Wang Y,
– review & editing. Petra Netter: Writing – original draft,

Smeland OB, Schork A, Holland D, Kauppi K et al.: Genome-wide
Writing – review & editing. analyses for personality traits identify six genomic loci and
show correlations with psychiatric disorders. Nat Genet 2017,
49
Important GWAS study investigating personality.
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