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Editorial on Individual Differences: from Neurobiological Bases to New Insight on Approach and Avoidance Behavior
The superordinate division of emotions is distributed along a bipolar dimension of affective valence, from approaching rewarding situations to avoiding punitive situations. Avoiding and approaching behaviors determine the disposition to the primary emotions of fear and attachment and the behavioral responses to the environmental stimuli of danger, novelty and reward. Approach or avoidance behaviors are associated with the brain pathways controlling cognitive and attentional function, reward sensitivity and emotional expression, involving prefrontal cortex, amygdala, striatum and cerebellum. Individual differences in approach and avoidance behavior might be modulated by normal variance in the level of functioning of different neurotransmitter systems, such as dopaminergic, serotoninergic, noradrenergic and endocannabinoid systems as well as many peptides such as corticotropin releasing hormone. These substances act at various central target areas to increase intensity of appetitive or defensive motivation. Physiologically, personality temperaments of approach and avoidance are viewed as instigators of propensity. They produce immediate affective, cognitive and behavioral inclinations in response to stimuli and orient individuals across domains and situations in a consistent fashion. Although the action undoubtedly emerges directly from these temperamental proclivities, ultimate behavioral outcomes are often a function of the integration among goal pursuit, self-regulation, and temperament trait. Defective coping strategies to aversive or rewarding stimuli characterize the patho-physiology of anxiety- and stress-related disorders or compulsive and addiction behaviors, respectively. Individuals with neuropsychiatric symptoms such as depression, suicidal behavior, bipolar mania, schizophrenia, substance use disorders, pathological gambling and anxiety disorders have scores which fall at the extreme tails of the normal distribution for a specific temperamental trait. The present Research Topic on the individual differences in emotional and motivational processing emphasizes the link between neuronal pattern and behavioral expression. The Topic includes experimental and clinical researches addressing the individual differences related to approach and avoidance and their behavioral characterization, structural and neurochemical profiles, synaptic connections, and receptor expressions. Studies are organized in a framework that puts in evidence the phenotypic expression and neurobiological patterns characterizing the individual differences and their biological variance
The role of neural and psychological factors in functional recovery
Recovery from brain insults which look similar at first glance may be a heterogeneous
phenomenon, since injury characteristics and neuropathological correlates
do not completely account for the variance in clinical outcomes, so that
predicting which patients will struggle and which will thrive is a difficult challenge.
The dissimilar behavioural and emotional outcomes often represent the
biggest barrier to rehabilitative treatment both in the acute phase and in social,
professional, and familial reintegration on the long term. Growing data indicate
that among the multiple neurobiological and psychological factors that impact on
the degree of brain recovery (Petrosini, 2017) there are the elements related to
pre-morbid personality and environmental context of the patient
Alexithymia: From neurobiological basis to clinical implications
Alexithymia is a construct of personality characterized by impairment in cognitive,
emotional and affective processing. It describes people with deficiencies in identifying,
processing, or describing subjective feelings or emotions. Although alexithymia is not a
psychological disorder per se and it is normally distributed in healthy population, it is
associated with enhanced risk of psychological impairment and it is present in a broad
spectrum of psychiatric and psychosomatic disorders, as chronic pain, somatoform
disorders, addictive disorders, anxiety and depression. By using neuroimaging studies,
variations associated with functional and structural differences in people with high
alexithymic traits are described in most of brain areas related to emotional awareness, as
anterior cingulate cortex, fusiform gyrus, amygdala, parahippocampal gyrus and insula.
Further, in the presence of alexithymia alterations are evidenced in the somato-sensory
and sensorimotor cortices as well as in cerebellar areas. The link between limbic and
somato-sensory systems may represent the possible neuroanatomical correlate of
alexithymia. In the present chapter, the neurobiological basis and clinical implications of
alexithymia are examined to clarify how the altered cognitive and affective experience of
emotion may result in a deficit in the emotional awareness
Cerebellum and personality traits.
Personality traits are multidimensional traits comprising cognitive, emotional, and behavioral characteristics, and a wide array of cerebral structures mediate individual variability. Differences in personality traits covary with brain morphometry in specific brain regions. A cerebellar role in emotional and affective processing and on personality characteristics has been suggested. In a large sample of healthy subjects of both sexes and differently aged, the macro- and micro-structural variations of the cerebellum were correlated with the scores obtained in the Temperament and Character Inventory (TCI) by Cloninger. Cerebellar volumes were associated positively with Novelty Seeking scores and negatively with Harm Avoidance scores. Given the cerebellar contribution in personality traits and emotional processing, we investigated the cerebellar involvement even in alexithymia, construct of personality characterized by impairment in cognitive, emotional, and affective processing. Interestingly, the subjects with high alexithymic traits had larger volumes in the bilateral Crus 1. The cerebellar substrate for some personality dimensions extends the relationship between personality and brain areas to a structure up to now thought to be involved mainly in motor and cognitive functions, much less in emotional processes and even less in personality individual differences. The enlarged volumes of Crus 1 in novelty seekers and alexithymics support the tendency to action featuring both personality constructs. In fact, Novelty Seeking and alexithymia are rooted in behavior and inescapably have a strong action component, resulting in stronger responses in the structures more focused on action and embodiment, as the cerebellum is
Cerebellum, Embodied Emotions, and Psychological Traits
This chapter addresses how the embodiment approach may represent a unifying perspective for examining the cerebellar role in emotional behavior and psychological traits. It is not intended to be exhaustive, but rather it can be a good starting point for advancing the cerebellar neural mechanism underlying embodiment. Our goal is to provide illustrative examples of embodied emotions and psychological traits in the emerging field of emotional and cognitive cerebellum. We illustrate how the cerebellum could be an important hub in the embodiment processes, associated with empathic abilities, impaired emotional identification and expression (as occurring for example in the presence of alexithymia), and specific psychological constructs (i.e., hypnotizability).This chapter addresses how the embodiment approach may represent a unifying perspective for examining the cerebellar role in emotional behavior and psychological traits. It is not intended to be exhaustive, but rather it can be a good starting point for advancing the cerebellar neural mechanism underlying embodiment. Our goal is to provide illustrative examples of embodied emotions and psychological traits in the emerging field of emotional and cognitive cerebellum. We illustrate how the cerebellum could be an important hub in the embodiment processes, associated with empathic abilities, impaired emotional identification and expression (as occurring for example in the presence of alexithymia), and specific psychological constructs (i.e., hypnotizability)
Viewing the personality traits through a cerebellar lens. A focus on the constructs of novelty seeking, harm avoidance, and alexithymia
The variance in the range of personality trait expression appears to be linked to structural variance in specific brain regions. In evidencing associations between personality factors and neurobiological measures, it seems evident that the cerebellum has not been up to now thought as having a key role in personality. This paper will review the most recent structural and functional neuroimaging literature that engages the cerebellum in personality traits, as novelty seeking and harm avoidance, and it will discuss the findings in the context of contemporary theories of affective and cognitive cerebellar function. By using region of interest (ROI)- and voxel-based approaches, we recently evidenced that the cerebellar volumes correlate positively with novelty seeking scores and negatively with harm avoidance scores. Subjects who search for new situations as a novelty seeker does (and a harm avoiding does not do) show a different engagement of their cerebellar circuitries in order to rapidly adapt to changing environments. The emerging model of cerebellar functionality may explain how the cerebellar abilities in planning, controlling, and putting into action the behavior are associated to normal or abnormal personality constructs. In this framework, it is worth reporting that increased cerebellar volumes are even associated with high scores in alexithymia, construct of personality characterized by impairment in cognitive, emotional, and affective processing. On such a basis, it seems necessary to go over the traditional cortico-centric view of personality constructs and to address the function of the cerebellar system in sustaining aspects of motivational network that characterizes the different temperamental trait
Trauma-related disorders and the bodily self: current perspectives and future directions
Trauma-related disorders are debilitating psychiatric conditions that influence people who have directly or indirectly witnessed adversities. Dramatic brain/body transformations and altered person's relationship with self, others, and the world occur when experiencing multiple types of traumas. In turn, these unfortunate modifications may contribute to predisposition to trauma-related vulnerability conditions, such as externalizing (aggression, delinquency, and conduct disorders) problems. This mini-review analyzes the relations between traumatic experiences (encoded as implicit and embodied procedural memories) and bodily self, sense of safety for the own body, and relationship with others, also in the presence of externalizing conducts. Furthermore, an emerging research area is also considered, highlighting principles and techniques of body-oriented and sensorimotor therapies designed to remodel bodily self-aspects in the presence of trauma, discussing their potential application with individuals showing externalizing problems
Individual differences in response to positive and negative stimuli: endocannabinoid-based insight on approach and avoidance behaviors
Approach and avoidance behaviors - the primary responses to the environmental stimuli of danger, novelty and reward - are associated with the brain structures that mediate cognitive functionality, reward sensitivity and emotional expression. Individual differences in approach and avoidance behaviors are modulated by the functioning of amygdaloid-hypothalamic-striatal and striatal-cerebellar networks implicated in action and reaction to salient stimuli. The nodes of these networks are strongly interconnected and by acting on them the endocannabinoid and dopaminergic systems increase the intensity of appetitive or defensive motivation. This review analyzes the approach and avoidance behaviors in humans and rodents, addresses neurobiological and neurochemical aspects of these behaviors, and proposes a possible synaptic plasticity mechanism, related to endocannabinoid-dependent long-term potentiation and depression that allows responding to salient positive and negative stimuli
From animal to human epigenetics
A rat story: Behavioral epigenetics beginnings
Human neurodevelopment is a dynamic and protracted process. It starts in the pre-natal life,
driven by genetic information, and continues unfolding following region-specific pathways up
to early adulthood (Gogtay et al., 2004; Koenderink & Uylings, 1995; Petanjek et al., 2011). Es
pecially during the pre-natal and early post-natal life, the developing brain depends on and is
sensitive to external inputs that shape its architecture and fine-tune neural connectivity patterns
according to environmental requirements (Branchi & Cirulli, 2014; Fox, Levitt, & Nelson, 2010;
Hensch, 2005; Takesian & Hensch, 2013; Teicher, Samson, Anderson, & Ohashi, 2016). Environ
mental inputs are therefore critical for a normative development. On the other hand, adverse
conditions occurring during sensitive periods for the nervous system maturation can interact
with genetic make-up and bias developmental trajectories toward maladaptive outcomes, as dem
onstrated by increased occurrence of psychopathology and psychiatric conditions following
childhood neglect, maltreatment and abuse (Benjet, Borges, & Medina-Mora, 2010; Bick &
Nelson, 2016; Cohen, Brown, & Smaile, 2001; Green et al., 2010; Kessler et al., 2010;
Widom, 1999).
It has been suggested that modifications of adult brain function and behavior changes induced
by early experiences can be determined by changes in the epigenetic status of specific genes (Bale
et al., 2010; Fraga et al., 2005; Maccari, Krugers, Morley-Fletcher, Szyf, & Brunton, 2014;
Tsankova, Renthal, Kumar, & Nestler, 2007). In fact, epigenetic mechanisms, including DNA
methylation, histone modifications, and non-coding RNAs regulation can be affected by various
extrinsic factors, providing a molecular link between external cues and gene expression (Kang
et al., 2011; Maze et al., 2014; Nord, Pattabiraman, Visel, & Rubenstein, 2015; Shibata,
Gulden, & Sestan, 2015).
Human studies evidence that individuals exposed to adversity in early post-natal life (Romens,
McDonald, Svaren, & Pollak, 2015; Tyrka, Price, Marsit, Walters, & Carpenter, 2012; van der
Knaap et al., 2014), or during pre-natal life (Mulligan, D’Errico, Stees, & Hughes, 2012; Perroud
et al., 2014) exhibit altered methylation of genes involved in hypothalamic-pituitary-adrenal (HPA)
axis functionality, as the NR3C1 gene coding glucocorticoid receptors (GR), a key element for the ho
meostasis of stress response system (Herman et al., 2016; Sapolsky, Meaney, & McEwen, 1985).
In turn, altered NR3C1 methylation levels have been associated to emotional and behavioral problems, externalizing and internalizing symptoms (Cicchetti & Handley, 2017; Dadds, Moul, Hawes, Mendoza
Diaz, & Brennan, 2015; Parade et al., 2016; Perroud et al., 2014; van der Knaap, van Oort, Verhulst,
Oldehinkel, & Riese, 2015). Further, reduced levels of NR3C1 messenger RNA (mRNA) and mRNA
transcripts, as well as increased cytosine methylation of the NR3C1 promoter were found in suicide
victims with a history of childhood abuse compared to suicide victims without childhood trauma
and controls (McGowan et al., 2009).
Finally, longitudinal studies on very preterm infant admitted to neonatal intensive care unit, and
thus subjected to pain-related stress and maternal separation, evidence an altered serotonin transporter
gene (SLC6A4) methylation status, predictive of enhanced socio-emotional stress reactivity and asso
ciated with less-than-optimal score at Personal-Social scale of Griffith Mental Development Scales at
12 months of age (Fumagalli et al., 2018; Montirosso et al., 2016; Provenzi, Guida, &
Montirosso, 2018).
Taken together, these findings corroborate an association between environmental experiences,
epigenetic modifications and behavioral outcomes.
Nevertheless, the cascade of biochemical events through which the environment is embedded in the
individual biology, affecting physiology and behavior remains unclear. In addition, not all individuals
exposed to early life adversity develop health issues, psychopathology or psychiatric disorders
(Collishaw et al., 2007; Yehuda & LeDoux, 2007). Though, the genetic make-up, epigenetic charac
teristics, and risk and protective factors that render individuals differently sensitive to environmental
influences are not yet understood (Belsky et al., 2009; Belsky & Pluess, 2009; Branchi, 2011).
Several aspects hamper the possibility to draw firm conclusions from human studies. Retrospective
designs rely on indirect information about the conditions of the participant, and even when information
is available or directly collected within prospective studies, it is virtually impossible to disentangle the
contribution of multiple factors occurring in pre-natal and post-natal life on specific physiological and
behavioral outcomes.
Moreover, both retrospective and prospective human studies depend on availability and access to
appropriate tissues for epigenetic analysis and are based primarily on saliva, blood and buccal cells
samples. Nonetheless, epigenetic patterns appear to be tissue and gene specific (Forest et al., 2018;
Smith et al., 2015) and there is little consensus on how much changes observed in peripheral tissues
may correlates each other and resemble changes in nervous tissue (Di Sante et al., 2018; Thompson
et al., 2013; Walton et al., 2016).
Animal models have strongly stimulated and complemented human studies (Phillips & Roth,
2019)(Box 1). Indeed, the animal models allow to prospectively manipulate the onset, quality, du
ration and predictability of environmental exposures under controlled conditions and to evaluate im
mediate, long-term and trans-generational consequences on candidate gene expression and behavior.
Laboratory animals can be exposed to aversive or permissive environments at different develop
mental time points and both genomic and non-genomic inheritance can be systematically investigated
(Bohacek & Mansuy, 2015, 2017; Francis, Diorio, Liu, & Meaney, 1999; Jirtle & Skinner, 2007;
Mitchell et al., 2016; Richards, 2006). In addition, since epigenetic reactions are bidirectional and po
tentially reversible (Cervoni & Szyf, 2001; Roth, Denu, & Allis, 2001) the causal relationship between
different epigenetic identities and behavioral outcomes can be addressed in animal models by admin
istering specific molecular compounds able to promote or inhibit epigenetic mechanisms as DNA
methylation (Keller, Doherty, & Roth, 2018, 2019; Weaver et al., 2004, 2005; Weaver, Meaney, &
Szyf, 2006).
Here, we report the contribution of animal studies, in particular, laboratory rats (Rattus norvegicus)
and mice (Mus musculus), to the field of human behavioral epigenetics. The chapter focuses on the role
of maternal environment as one of the most studied vectors in inducing epigenetic modifications and
enduring phenotype in the offspring. In addition, studies on the interplay among genetic polymor
phisms, epigenome and aversive environments in contributing to psychiatric disorders and, as a bright
note, on the effect of the exposure to permissive environment (with attention to environmental enrich
ment) will be reported
Features of sequential learning in hemicerebellectomized rats
Because the sequencing property is one of the functions in which cerebellar circuits are involved, it is important to analyze the features of sequential learning in the presence of cerebellar damage. Hemicerebellectomized and control rats were tested in a four-choice visuomotor learning task that required both the detection of a specific sequence of correct choices and the acquisition of procedural rules about how to perform the task. The findings indicate that the presence of the hemicerebellectomy did not affect the first phases of detection and acquisition of the sequential visuomotor task, delayed but did not prevent the learning of the sequential task, slowed down speed-up and proceduralization phases, and loosened the reward-response associative structure. The performances of hemicerebellectomized animals in the serial learning task as well as in the open field task demonstrated that the delayed sequential learning task could not be ascribed to impairment of motor functions or discriminative abilities or to low levels of motivation. The delay in sequential learning observed in the presence of a cerebellar lesion appeared to be related mainly to a delay of the automatization of the response. In conclusion, it may be advanced that, through cortical and subcortical connections, the cerebellum provides the acquisition of rapid and accurate sensory-guided sequence of responses. (C) 2009 Wiley-Liss, Inc
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