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Neural Control of Behavior

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N
Neural Control of Behavior
Ashutosh Kumar1, Ravi Kant Narayan1,
Vikas Pareek2, Chiman Kumari4, Sanjib K. Ghosh1
and Muneeb A. Faiq3
1
Department of Anatomy, All India Institute of
Medical Sciences (AIIMS), Patna, India
2
Computational Neuroscience and Neuroimaging
Division, National Brain Research Centre
(NBRC), Manesar, India
3
Neuroimaging and Visual Science Laboratory,
Langone Medical Centre, New York University
School of Medicine, New York, NY, USA
4
Department of Anatomy, Postgraduate Institute
of Medical Education and Research (PGIMER),
Chandigarh, India
Introduction
The way in which an individual acts in response to
a stimulus or situation is called “behavior.”
Behavior is chiefly controlled by the central ner-
vous system (i.e., brain and spinal cord). Behavior
can be innate (present since birth) or is learned
during a lifetime. Further it can be simple or
complex. The simplest forms of the behavior are
the “reflexes” which are innate in nature. For a
reflexive behavior, the individuals may fail to
perceive a conscious appraisal, as it primarily
gets executed from the motor centers in the brain
stem or spinal cord. Complex behavior, a varying
motor action or response adjusted to the need of a
© Springer Nature Switzerland AG 2019
J. Vonk, T. K. Shackelford (eds.), Encyclopedia of Animal Cognition and Behavior,
https://doi.org/10.1007/978-3-319-47829-6_1368-1
situation, involves more than one component of
the brain and is intricately regulated. A stimulus
which is received at the sensory receptors is car-
ried by the nerves to the sensory cortex which
generates perception and further sends it to the
cognitive and the effectors domains of the brain
where complex neural processing constructs a
behavior from the substrate, in response. The
comprehensive cortical processing of sensory
feedback is essential for the conscious appraisal
of a complex behavior. Reflex behavior is com-
mon to all the organisms, while complex behav-
iors are unique to the animal kingdom and denote
their intellectual ability and hierarchy in the living
system.
Complex behaviors can be innate or learned.
Innate behaviors (as feeding or sexual behavior)
are genetically determined. Innate behaviors
involve hypothalamus and other subcortical cen-
ters and are associated with limited cortical pro-
cessing in comparison with the learned behaviors.
Emotional behaviors, which are largely innate, are
regulated by limbic system which presents a
unique set of cortical and subcortical centers.
Learned behaviors may be driven by reward – a
phenomenon called conditioning – or may not
involve any reward at all, called nonassociative
learning. A reward pathway which connects
aminergic nuclei of the brain stem to the specific
nuclei group in the forebrain has been considered
as the driver of any reward-based behavior
(Kiernan et al. 2014). Aminergic nuclei in the
brain stem are known as the prime determinants
2 Neural Control of Behavior
of the adaptive behavior – which is largely a
learned behavior helping the individuals in
adapting to the environmental challenges.
How brain generates behavior has been an all-
time curiosity of the human which has now devel-
oped into a distinct stream of the neuroscience,
called neuroethology. In recent years, knowledge
of the neural control of the behavior has got
extensive research updates revolutionizing the
existing understanding of the neuroethology.
Behavioral roles have been found for many new
brain regions which were earlier either not or little
known. Recent research suggests extensive role of
anterior cingulate cortex in decision-making.
Basal nuclei and cerebellum have been found to
bear important roles in behavior, in contrast to
their earlier known roles which limited to the
movement control. There are also updates regard-
ing the role of the insula (a cortical island) and
claustrum (a strip of gray matter), for which any
cognitive/behavioral functions were earlier little
known. Uniquely, habenula – a subcortical set of
nuclei which connects with brain stem – has been
noted crucial in regulating adaptive behavior
(Kumar et al. 2017b). Other than the central ner-
vous system, two other nervous system types, i.e.,
“autonomic” and “enteric,” are also known to bear
important roles in complex behavior. The current
research adds extensively into their behavioral
functions.
Present chapter takes a comprehensive look on
the role of the three nervous system types
involved in regulation of the animal behavior,
and behavior-specific neural circuitries comprised
of them, with a focus in human.
Methods for Study of Neural Regulation
of Behavior
Animal behavior is either studied by direct obser-
vations of its response to a given stimulus in a
natural setting or through modeling of a behavior
in laboratory settings. In laboratory settings, inter-
ventions like creating a lesion or inserting an
implant, or the introduction of a pharmacological
agent into the nervous system of an animal, can be
done to model a behavior. These models can be
further assessed by the standard neuropsycholog-
ical tests.
For long, creating a lesion in the brain to know
its impact remained a primary method to study a
behavior. Implying the electrodes in targeted brain
parts to record the local field potential (LFP – an
electrophysiological signal created by summed up
electric current arising from large population of
neurons from the recording site) in the behavior-
ally activated neurons or to observe effect of an
electrical stimulation, or pharmacological inter-
ventions, had been the next prevailed methods to
study animal behavior in laboratory settings.
These methods when combined with the neuro-
imaging modalities have brought revolution in the
understanding of neural control of behavior.
There have been significant advances in neu-
roimaging (and recording) methods in recent
years which have made structure-function cou-
pling for the brain parts possible. Multimodal
neuroimaging and data integration from chosen
combinations of the magnetic resonance imaging
(MRI), positron emission tomography (PET),
single-photon emission computed tomography
(SPECT), near-infrared spectroscopy (NIRS),
magnetoencephalography (MEG), electroenceph-
alography (EEG), diffuse optical tomography
(DOT), etc. have given edge to the behavioral
research. The advances have given rise to a new
stream of behavioral study, neuroeconomics – an
interdisciplinary field which seeks to explain how
decisions are made in the brain (Xue et al. 2010).
Structural magnetic resonance imaging (sMRI)
has now got resolution of 7 and 9 tesla which is
able to provide the structural details of the brain
parts in living closer to that histological study in
postmortem brain tissue. Use of contrast-based
sMRI (mainly iron oxide) can detect the activity
of targeted molecules with high sensitivity and
specificity.
Diffusion tensor imaging (DTI) has revolution-
ized the knowledge on the fiber connections of the
brain parts. DTI is a sub-modality of the structural
MRI that encodes diffusion effects of the water
molecules in the nuclear magnetic resonance sig-
nal by using bipolar magnetic field gradient
pulses. The fiber tracts can be delineated in vivo
based on the fractional anisotropy. Constrained
Neural Control of Behavior
spherical deconvolution (CSD) tractography is a
modification of DTI which can provide better
three-dimensional model of the neural tracts due
to its ability to detect crossing fibers in the tracts.
Functional magnetic resonance imaging
(fMRI) can depict brain activity by detecting the
associated changes in brain hemodynamics by
using blood-oxygen-level-dependent (BOLD)
contrast. BOLD reflects cerebral blood flow
(CBF) as brain function requires blood flow to
supply oxygen for energy consumption by neu-
rons. Event-related fMRI can also be used to label
specific molecules with contrast agents called
molecular MRI. A statistical method called
multi-voxel pattern analysis (MVPA) implies feed-
ing of fMRI data into a computer algorithm that
automatically learns the neural patterns associated
with specific thoughts or experiences.
Single-photon emission computed tomography
(SPECT) is a functional nuclear imaging method
often used to image awake behaving animals
(where a targeted brain part is stimulated using
invasive electrodes or some other physical
means). It uses an intravenous radiotracer to
detect cerebral perfusion during activity and in
turn neuronal activity (which depends on cerebral
perfusion rate).
Positron emission tomography (PET) is the
best to study neurotransmitter/receptor interac-
tions. It has lower spatiotemporal resolution
when compared to MRI. It is exquisitely sensitive
for detecting the targeted molecules or processes.
EEG and MEG detect synchronized activity of
an assembly of neurons by measuring LFP or
magnetic fluxes. EEG and MEG can be used to
explore brain cortical activation pattern with ultra-
high temporal resolution and real-time recording
of the event-evoked neural information flow.
Stereoelectroencephalography (SEEG) uses
in-depth electrodes (surgically implanted in the
brain tissue) for recording electroencephalo-
graphic signals from desired subcortical nuclei in
animal models.
Electrical recording of the cortex and in-depth
brain parts has progressed much in recent years.
Scalp electrodes have become more efficient. For
intracranial recording, miniature multiplex elec-
trode panels or multielectrode arrays (MEAs) can
3
get surface recordings from large parts of the
cortex very easily, and new hair-thin electrodes
can reach in-depth brain parts without causing any
significant brain damage. Wireless electrodes can
be implanted in brain parts in behaving animals
which can be operated from remote control to
provide stimulation or recording. Magnetic trans-
cranial and deep brain stimulation are now being
frequently used to stimulate desired brain parts.
Whole cell patch clamp (which can detect ionic
currents in single neurons) can be used to detect
neuronal activity at the level of single neurons in
awake or behaving animals. Whole brain imaging
of neuronal activity using fluorescent calcium
indicators and multiphoton laser scanning micro-
scopes in head-fixed behaving animals is a recent
method for detecting behavior-specific neuronal
circuitry.
Diffuse optical tomography (DOT) is an opti-
cal spectroscopy method which uses infrared light
to probe activity inside any organ. Light entering
in the tissue can undergo multiple scattering, and
as a result some photons are redirected back
toward the tissue surface each time. The distance
this light travels and the changes in its physical
properties in that course carry information which
can be decoded to measure events and map their
locations within the brain tissue. Regional varia-
tions in oxy- and deoxy-hemoglobin concentrations
as well as blood flow and oxygen consumption can
be imaged by monitoring spatiotemporal varia-
tions in the absorption spectra of used light.
DOT can be used to monitor brain activity; it can
directly measure hemodynamic, metabolic, and
neuronal responses to neurons and tissue and
organ activations.
Newer Methods
Use of inducible gene constructs for the mem-
brane ion channels in model animals can provide
high-quality information for the neural circuitry –
behavior relationship. Chemogenetics uses system-
atically administered Designer Receptors Exclu-
sively Activated by Designer Drugs (DREADD)
targeted to the specific molecules (Kumar et al.
2018c). Optogenetics uses laser lights of specific
wavelengths to control activation of ion channels
in neurons that are genetically tagged with
4 Neural Control of Behavior
fluorescent biomolecules. Fine optical electrodes
are inserted in the brain for light-induced manip-
ulation of electrical activity in in-depth neurons
and neuronal circuits. This method has revolution-
ized the study of animal behavior (Deisseroth
2010; Kumar et al. 2018a). Sonogenetics uses
low-pressure ultrasound waves to activate the arti-
ficially ultrasonically sensitized neurons (Ibsen
et al. 2015). Brain-computer interface (BCI) inte-
grated with the neuro-recording methods like
EEG is being used as a mode of study or manip-
ulating a specific behavior based on the
neurofeedback (Grau et al. 2014; Kumar et al.
2018a).
Computational modeling (CM) is a recent
entry into the study of animal behavior. It uses
computers to simulate and study the behavior of
complex systems applying principles of mathe-
matics, physics, and computer science. The bene-
fit of using this method is that it can predict the
influence of numerous variables which character-
ize any system, in minuscule time, with minimum
of efforts by creating simulation of the system
(Pareek 2017). A simulation is created by manip-
ulating the variables individually or in combina-
tion and observing how the alterations influence
the outcomes – called in silico experiment. Com-
putational models have the ability to study a sys-
tem at multiple levels – known as multiscale
modeling (Cipresso 2015).
CM is a unique tool to understand how indi-
viduals or groups influence each other’s behavior
in a social structure. Virtual reality (VR)-based
experimental settings are used to manipulate
behavior in a single or a small group of individ-
uals, and information-based parameters are extra-
cted to develop a basic computational model of
the behavior dynamics. (VR is a computer-
generated simulation of the reality.) These basic
models are further used to develop simulations to
understand and explain behavior dynamics at the
social level in the context of the possible scenar-
ios. CM can also be used to study behavioral
pathologies in individuals or groups (Cipresso
2015).
Neurodevelopmental Basis of Behavior
The earliest forms of behaviors in a growing fetus
are simple sensorimotor reflexes when brain parts
and neuronal tracts are still maturing. The primi-
tive reflexes disappear after birth. Any return of
the primitive reflexes in an adult is considered as
the indicator of a neuronal lesion. Childhood and
adolescent ages are the critical periods when an
individual forms concepts about the environment
surrounding it and develops “theory of mind, an
ability to understand mental status of others,”
which are very quintessential for learning social
behaviors. As cognitive and behavioral domains
are still maturing, children and adolescents are
vulnerable to develop behavioral disorders. Poor
rearing up conditions during childhood and ado-
lescence are known to adversely affect social
behavior of the individual. Certain individuals
are considered more susceptible to develop such
behavioral disorders due to their unique genotypic
structure.
Common Neural Mechanisms in
Complex Behaviors
Perception-Cognition-Action Triad
A behavior generates from the hierarchically
arranged cortico-brain stem/spinal networks
(top-down and bottom-up) of the central nervous
system (CNS). Perception and cognition are the
neural substrates for any behavior. Any given
stimulus is received by a peripheral sensory
organ which is further carried to sensory cortex
of the brain through sensory pathways (entering
either through the spinal cord or brain stem at the
lowest hierarchical level and then through the
midbrain and the diencephalon). In the sensory
cortex, perception for the stimulus is generated
(which can be further unified in a multimodal
cortical area with the corroboratory perceptions
from other sensory modalities). The perception is
further encoded by cognitive domains of the brain
in the form of a memory or learning, following
which effectors domains generate a suitable
response for the encountered stimulus based on
the merits.
Neural Control of Behavior
Prediction Signaling, Error, and Response
It is a basic property of any neuronal ensemble
that it would generate a prediction estimate
against any incoming sensation or a motor com-
mand. This prediction (or prediction signaling) is
further matched with the actual input to the nuclei.
Any difference of the actual and expectant value is
called prediction error which is used to correct the
future predictions. Neuronal ensembles calculate
saliency (strength) and valence (positive or nega-
tive character, which is important in reward pre-
dictions) of a stimulus. Any behavioral response
is based on the prediction error values (den Ouden
et al. 2012). Any behavioral response against a
novel stimulus generates learning which can be
stored in the specific parts of the brain as the
memory patterns (or engrams), which will provide
an experienced-based contribution to the future
behaviors when a similar stimulus is encountered.
A prediction error-based construction of the
behavior will be further discussed in the section
“Behavior-Specific Neural Correlates” of this
entry.
Behavior-Induced Brain Plasticity and Vice
Versa
Behavior induces plastic changes in the brain
components. The plasticity may involve brain
part (like size of amygdala may increase in the
anxiety disorders), connections, dendritic arbori-
zation and spine density, axonal sprouting, synap-
tic organization and formation, and neurogenesis
(in hippocampus). Vice versa, the brain plasticity
may have influence on cognitive abilities and
behavior of the individual which in turn may
induce further plasticity – a phenomenon known
as meta-plasticity (Altenmüller and Furuya
2016) – and is regarded crucial for the behavioral
modifications and learning motor skills.
Role of Neurotransmitters, Neuromodulators,
and Neurohormones
Neurotransmitters are the chemical messengers
which facilitate communication of impulses at syn-
apses. Neuromodulators (catecholamines deriva-
tives such as dopamine, serotonin, noradrenaline,
and many small peptides like enkephalin, endor-
phin, dynorphin, etc.) and neurohormones (which
5
are secreted by neurons, like estrogen, oxytocin,
and vasopressin) have the capacity to modulate
transmission at the synapses. Though a behavior
is not directly created at the synapses, synaptic
communications create neural substrate for it.
Along with the neurotransmitters, neuromodulators
and neurohormones have critical roles in setting
and maintenance of the mood. An intricately reg-
ulated cocktail of these neurochemicals not only
determines the quality of synaptic communication
but also sets the rate, rhythm, coupling, and syn-
chronization of the oscillatory waves arising from
cortical neurons – which is crucial for the con-
struction of an appropriate response to the
encountered stimulus (Kumar et al. 2018b).
Neural Basis of Sexually Dimorphic Behaviors
We can distinguish the behavioral differences
between male and female by looking at their vary-
ing responses to the same environmental stimuli,
but neuroscientific basis of this is not very clear
yet. The new research has gradually disapproved
the earlier belief that some brain parts are sexually
dimorphic. Accumulating evidence suggests the
sex-specific differences are not in the gross brain
structures or neural connections but lie at the
ultrastructural levels like transcriptomic and pro-
teomic configuration of the specific brain parts
and molecular arrangement and signaling at the
neuronal synapses (Chen et al. 2019).
Salient Brain Parts/Nervous System
Components and their Affiliated
Behaviors
Cerebral Cortex: Supreme Controller of
Animal Behavior
Ordinarily, the cerebral cortex is believed to have
control over cranial and spinal regulation of motor
actions. It also has limited control over basic/
instinctive behaviors as hunger, thirst, or sex,
which primarily involve hypothalamus. Brain
has primarily four cortical lobes (frontal, parietal,
temporal, and occipital lobes) controlling all kinds
of behavior. Two minor lobes – insular and
limbic – are also described which involve parts
of the four primary lobes. (A detailed description
6 Neural Control of Behavior
on behavioral role of insular cortex has been pro-
vided later in this entry in segment “Emerging
behavioral role for other brain parts and nervous
system components.” Limbic cortex is further
described with limbic system in section “Emo-
tional Behavior” under section “Behavior-
Specific Neural Correlates.”) Prefrontal cortex
(PFC) (anterior part of the frontal lobe) is the
most important cortical part involved in the con-
trol of executive, social, emotional, or instinctive
behaviors. Distinctive parts of the prefrontal cor-
tex are involved in specific aspects of the behav-
ior, e.g., dorsolateral prefrontal cortex (DLPFC) is
involved in executive behaviors (showing intelli-
gence) like planning, thinking, making judg-
ments, and problem-solving. Ventrolateral and
ventromedial/orbital (VLPFC and VMPFC or
OFC) parts concern more to the control of social
and emotional components of a behavior. Anterior
cingulate cortex (ACC) is said to have crucial role
in decision-making and personal/social conflict
resolution. VLFC, VMFC/OFC, and ACC are
also involved in fetching attention of the individ-
ual to a salient stimulus, hence working as a self-
referencing center.
Cortical Modularity-Based Cerebral
Mechanisms
Cortical domains in the brain are arranged into
large-scale structural and functional networks.
A large-scale network can be decomposed into
sets of densely connected small-world networks
(based on the short average distances or closeness
between connecting elements) and activated dur-
ing a specific cognitive function or sensory per-
ception (Meunier et al. 2010). Major cognitive
cortical modules are saliency network (SN),
default mode network (DMN), dorsal attention
network (DAN) and ventral attention network
(VAN), task-positive network (TPN), and central
executive network (CEN). Each such small-world
cortical network is defined as a module – which
can be represented on a graph and contains dense
structural/functional connections (called edges)
between its components (called nodes). A cortical
module is an evolutionarily conserved and func-
tionally segregated network. There could be fur-
ther levels of hierarchy within a module, where it
can be decomposed into more densely connected
clusters or subsets (Meunier et al. 2010). These
cortical modules are delineated by the graph-
theoretic study of fMRI connectivity pattern dur-
ing rest- and task-related activity. DMN and exec-
utive/attention networks exist in anticorrelation
(activation of one has deactivating influence on
another) relationship and deal with “task-negative”
and “task-positive” cognition, respectively. DMN
gets activated when an individual is at rest,
absorbed into autobiographical moments, or self-
monitoring detached from the external world.
Conversely, executive/attention networks get acti-
vated when individual follows a task or engages to
the external world. The DMN grossly includes
VMPFC, bilateral posterior cingulate cortex
(PCC) and retrosplenial cortex (RspC), inferior
parietal lobule (IPL), parts of the hippocampal
formation, medial temporal lobes (MTL), and
the angular gyrus. The CEN is anchored in
DLPFC and PPC. The DAN is organized bilater-
ally and grossly comprises the intraparietal sulcus
(IPS) and the frontal eye fields (FEF) of each
hemisphere. The VAN which is said to be
lateralized to the right hemisphere involves
temporoparietal junction (TPJ) and the ventral
frontal cortex (VFC). Salient network (SN) com-
prises anterior insula (AI) and dorsal anterior cin-
gulate cortex (dACC). SN detects relevant internal
or external stimuli and also acts as a switch
between DMN and ACN (and DAN/VAN).
Other than above described major cortical
modules, there are many minor modules, like
that for face, object, color identification, or mem-
ory recall. Major sensory perception cortical mod-
ules are that for visual, olfactory, tactile, and
auditory perception.
Cortical modules hold key to specific neuro-
physiological processes or sensory perceptions,
and cognitive functions (Hilger et al. 2017), and
are building blocks for any behavior. Several of
the cortical modules can be recruited to orches-
trate behavioral response to any stimulus.
Theory of Mind: A Basic Cortical Function in
Social Behavior
Theory of mind is defined as a cognitive ability to
understand mental status of others and develops
Neural Control of Behavior
during childhood and adolescence (as we
described above). This ability is more evident in
human, primates, and other animals showing
higher engagement in social behavior. This is an
essential ability for peaceful coexistence in a soci-
ety which depends on strong social relationships
and cooperations with each other. Neural corre-
lates of theory of mind are not much understood
though mainly cortical parts (prefrontal cortex,
insular cortex, parts of parietal and temporal cor-
tices (inferior parietal lobule, temporoparietal
junction, posterior part of the superior temporal
sulcus, etc.)) are said to contribute to this.
A unique type of cortical neurons – “mirror neu-
rons which activate when animals view others
doing actions” (Carrillo et al. 2019) – is said to
be crucial in developing this ability (has been
described in detail along with “insula and mirror
neurons” in section “Emerging Behavioral Role
for Other Brain Parts and Nervous System Com-
ponents”). Another type of neurons (similar to the
mirror neurons in cortex) was recently reported in
amygdala, called simulation neurons, which help
in predicting and creating a mental simulation of
others’ probable actions based on their observa-
tions (Grabenhorst et al. 2019) (further studies
will be necessary to develop a concrete concept
in this regard).
Hypothalamus: Master Regulator of Innate
Behavior
Hypothalamus is also called as visceral brain due
to its concern with the visceral functions like
feeding, hunger, and thirst. It is also involved in
the instinctual behavior like sex and reproduction.
Hypothalamus is also the master regulator of
endocrine organs, and it is the highest autonomic
center in brain (Kumar et al. 2018c). Thus, hypo-
thalamus presents a hub connecting the endocrine
regulatory axis and autonomic nervous system to
the rest of the brain.
Hippocampus: A Memory Base for the
Cognitive Flexibility and Social Behavior
The role of hippocampus has been considered
important in creating cognitive flexibility and
social behavior (Rubin et al. 2014). Hippocampus
exerts its influence on behavior through memory.
7
In hippocampus, registration and long-term
potentiation (LTP) of the memory occur
(Kiernan et al. 2014). Memory contents create a
neural substrate for any experience-based behav-
ior. Hippocampus has an exclusive role in learn-
ing which can create new experiences which in
turn can amend an old behavior. Nearby para-
hippocampal region (entorhinal cortex) is known
to register spatiotemporal details of the surround-
ings the animal resides in that creates important
attribute to a learned behavior. Hippocampus has
also an influence on the hypothalamo-pituitary-
adrenal (HPA) axis, through which it regulates
endocrine homeostasis, which in turn can influ-
ence behavior.
Hippocampus Microanatomy (In Brief): The
hippocampus is located in the medial temporal
lobe. It contains two parts: the hippocampus
proper and the dentate gyrus. The hippocampus
and dentate gyrus (DG) have the shape of a curved
tube, which can be compared to a seahorse and a
ram’s horn (Cornu Ammonis – CA). Hippocam-
pal subfields CA1, CA2, and CA3 are named by
abbreviation of Cornu Ammonis. Hippocampus
proper and DG present an allocortex bearing
only three layers. The major neurons in the hip-
pocampus proper and DG are pyramidal neurons
and granule cells, respectively. The hippocampus
is reciprocally connected to various cortical and
subcortical structures. Its prime cortical connec-
tion is from the entorhinal cortex (EC). The EC is
located in the parahippocampal gyrus, a cortical
region adjacent to the hippocampus bearing all six
layers. The cortical transition zone (from three
layers to six layers) between the hippocampus
and EC is called subiculum. The entorhinal cortex
(EC) is strongly and reciprocally connected with
many cortical and subcortical structures as well as
with the brain stem. Its major efferents to the
hippocampus enter via perforant pathway
bypassing the subiculum (Kiernan et al. 2014).
Figure 1 provides a schematic description of the
chief hippocampal microcircuitry connecting to
EC. The reentrant signals through hippocampal-
EC loop are associated with registration and long-
term potentiation of the new memories.
8 Neural Control of Behavior
Neural Control of Behavior, Fig. 1 Hippocampal
microcircuitry in behavior (Sensory signals drive the
perforant path (PP, black) excitatory inputs from EC II
and III layer pyramidal neurons to granule cells in the
dentate gyrus (DG, black), which in turn sends mossy
fiber axons (black) to CA3, and then CA3 feeds excitatory
inputs onto CA1 neurons through Schaffer collaterals (SC,
Basal Nuclei: A Tool for Selection of
Appropriate Behavior
Basal nuclei (BN, earlier known as basal ganglia)
are a set of subcortical gray matter collections
located in vicinity of the diencephalon (Kiernan
et al. 2014). Earlier known roles of basal nuclei in
behavior were limited to the motor functions like
movement, but new research establishes its essen-
tial role in reward value-guided or motivated
behavior – it helps to decide which choice will
be more rewarding to follow (Hikosaka et al.
2014). Reward can be evaluated here for the
short-term and long-term benefits – which give
individual chance to switch from a choice based
on its current value (Kim and Hikosaka 2015).
Simply put, basal nuclei help to select most appro-
priate behavior among the available alternatives in
response of an external or internal stimulus. The
appropriateness of a choice is decided by compet-
itive actions between three pathways in the basal
nuclei neural circuitry, namely, direct, indirect,
and hyperdirect pathways (Nambu 2009).
black). A major output of the hippocampus arises from
CA1 pyramidal neurons, which projects back to EC
(V and VI layers) thus completing the hippocampal-EC
loop. Hippocampal microcircuitry also contains
GABAergic interneurons (red) which modulate pyramidal
neuron activity in a domain-specific manner)
Basal Nuclei: Basic Anatomy (In Brief)
The main components of the basal nuclei – as
defined functionally – are the striatum (dorsal
striatum comprising of caudate nucleus and puta-
men and ventral striatum comprising of nucleus
accumbens and olfactory tubercle), globus
pallidus, substantia nigra, and subthalamic nucleus
(STN). The globus pallidus is functionally
divided into external (GPe) and internal (GPi)
domains. The substantia nigra which is actually
a part of the midbrain is functionally divided into
two parts: the inner pars compacta (SNc) and the
outer pars reticularis (SNr). The basal nuclei pre-
sent a three-tier structure (input, caudate nucleus
and putamen; intrinsic, GPe, STN, SNc; and out-
put GPi, SNr nuclei) connecting cerebral cortex in
a cortico-basal nuclei-thalamo-cortical (CBnTC)
loop.
Basal nuclei microcircuitry in behavior is orga-
nized in CBnTC loops. There are four types of
CBnTC loops which can be recruited for any
behavior (Table 1) (Kiernan et al. 2014).
The three known basal nuclei pathways (direct,
indirect, and hyperdirect) involve one of these
Neural Control of Behavior 9

Neural Control of Behavior, Table 1 Cortico-basal nuclei-thalamo-cortical loops in behavior

Cortico-basal
nuclei-thalamo-
cortical (CBnTC) Part of Thalamic
loops Cortical input striatum Part of pallidum nucleus Cortical output
Motor Primary somatic Putamen Globus pallidus Ventral Supplementary and
sensory and motor lateral and primary motor areas
areas; premotor ventral and premotor area
area anterior
nuclei
Oculomotor Prefrontal and Caudate Globus Ventral Frontal eye fields
posterior parietal nucleus pallidus; anterior and
cortex substantia nigra mediodorsal
pars reticulata nuclei
Prefrontal Premotor area and Caudate Globus pallidus Ventral Prefrontal cortex
posterior parietal nucleus anterior and
cortex mediodorsal
nuclei
Limbic Temporal lobe; Nucleus Ventral Mediodorsal Cingulate gyrus and
hippocampal accumbens pallidum nucleus orbital prefrontal
formation; (ventral cortex
amygdala striatum)
Adapted from Bar’s The Human Nervous System: An Anatomical Viewpoint, Tenth Edition

CBnTC loops. Figure 2 presents a schematic dia- Cerebellar Microanatomy (In Brief)
gram of the basal nuclei microcircuitry implied in A histological section of cerebellum would show
behavior. an outer folded cortex (gray matter) surrounding
an inner medulla (white matter) which contains
deep nuclei embedded within it. There are three
Cerebellum: A Prime Moderator for all
layers to the cerebellar cortex; from outer to inner,
Behavioral Actions
these are the molecular, Purkinje, and granular
Like the basal nuclei, the cerebellum was primar-
layers. The function of the cerebellar cortex is
ily known for its role in movement. New evidence
essentially to modulate information flowing
suggests the cerebellum is involved in moderation
through the deep nuclei embedded in the medulla.
of each of the functions of the cerebrum; hence
The molecular layer is outermost layer of the
supposedly it has a function to modulate the cere-
cerebellar cortex, containing two types of inhibi-
bral control of cognition and behavior. It was
tory interneurons: the stellate and basket cells. It
reported to be involved in the control of the
also contains the dendritic arbors of Purkinje neu-
reward-oriented behavior mediated by its projec-
rons and parallel fiber tracts (axons of granule
tion to VTA – a brain stem dopaminergic nuclei
cells), coming from the granular layer. The middle
which also gets projections from the prefrontal
layer contains only one type of cell body – that of
cortex (Carta et al. 2019). It gets activated several
the large Purkinje cells. Purkinje cells are the
milliseconds prior to the cerebrum when we are to
primary integrative neurons of the cerebellar cor-
initiate an action or even in showing imaginary of
tex and provide its sole output which is inhibitory
an action (Kiernan et al. 2014). It assembles and
in nature. The innermost layer contains the cell
provides experience-based feedback for correc-
bodies of two types of cells: the numerous and
tion of the prediction error (made by executive
smaller granule cells and the larger Golgi cells
and motor centers in cerebral cortex) which brings
(Kiernan et al. 2014). Figure 3 provides a sche-
perfection to a learned behavior (Rapoport
matic description of functional cerebellar micro-
et al. 2000).
circuitry implied in behavior.
10
Cerebral
Cortex
Glutamate, ve+
Hyper-DIRECT
PATHWAY
Striatum
(CAUDATE + PUTAMEN)
GABA, ve.
In-DIRECT
Dopamine,
PATHWAY ve.via D2
Globus Pallidus
EXTERNAL
Less GABA
released,
less ve.
Glutamate released, ve+
Sub-thalamic nucleus
More Glutamate released, ve++
Hyper-DIRECT PATHWAY
In-DIRECT PATHWAY
DIRECT PATHWAY
Common transmissions in all the 3 pathways
Neural Control of Behavior, Fig. 2 Basal nuclei
microcircuitry in behavior (All the basal nuclei pathways
begin with the cerebral cortex stimulating striatum by the
glutamatergic efferents. Thereafter, striatum inhibits
Globus pallidus “externa” in “In-Direct” and “interna” in
“Direct” pathway by the GABAergic efferents. An
inhibited Globus pallidus “interna” releases less amount
of GABA to the adjacent Sub-thalamic nucleus, which is
directly stimulated by the efferents from cerebral cortex in
“Hyper- Direct” pathway. Subthalamic nucleus in turn
stimulates the “interna” by more amount of glutamate
Reticular Formation (RF): Center for the
Generation of Consciousness, Arousal, and
Behavioral Reflexes
RF is a diffuse neural network comprising of
neuronal cell bodies and nerve fibers located in
brain stem core. RF contains diverse set of nuclei
arranged in columns which send axons up and
down to almost all brain parts including cerebral
cortex and cerebellum and spinal cord segments.
Their dendrites create an entangled network
locally, contact with the cranial nerve nuclei
which are located in the brain stem, and receive
collaterals from ascending and descending tracts
Neural Control of Behavior
Selection of appropriate behaviour
depending on competing pathways
Glutamate, ve+
in In-Direct (ve.) & Hyper-direct pathway (ve..)
Less Excitatory signal
in Direct Pathway, more
Excitatory signal, ve+
Dopamine,
ve+ via
D1
GABA, ve.
Thalamus
(VA/VL nuclei)
Substantia
Nigra
Pars Compacta
DIRECT in In-Direct (ve.) & Hyper-direct pathway (ve..)
PATHWAY more GABA released
in Direct Pathway less GABA
released, less ve.
Globus Pallidus
INTERNAL
being released from it. Efferents from Globus pallidus
“interna” now inhibit the thalamus, the extent of which
depends upon the pathway being followed, more inhibition
in In-Direct and Hyper-direct pathway as compared to that
in Direct pathway. The thalamus by default stimulates the
cortex by the glutamatergic efferents, which is more in
Direct pathway (disinhibition) than the rest of the two
pathways (inhibition). Depending upon the resultant stim-
ulation of the motor cortex, appropriate behavior can
ensue. The inhibition made by Hyper-Direct pathway is
more than the In-Direct pathway)
passing through the brain stem (except that in the
posterior column for fine touch, pressure, and
proprioception) (Kiernan et al. 2014). Among
RF nuclei also are the aminergic nuclei which
secrete adrenaline/noradrenaline, dopamine sero-
tonin, acetylcholine, etc. and take part in reward-
oriented and adaptive behavior. Ascending corti-
cal projections from RF, called reticular activating
system (RAS), relay through nonspecific midline
nuclei of thalamus and then spread diffusely to all
over the cortex. RAS is said to provide arousal to
the cortex hence to contribute to the alertness and
attention. Secretions of aminergic nuclei which
Neural Control of Behavior
Neural Control of Behavior, Fig. 3 Cerebellar micro-
circuitry in behavior (The Climbing fibers and Mossy
fibers are the two afferent fibers to the cerebellum. The
Climbing fibers provide excitatory input from the inferior
olivary nucleus of the medulla, on dendritic shafts of the
Purkinje cells. The Mossy fibers are coming mainly from
the cerebral cortex via pontine nuclei. The Mossy fibers
synapse with the Granule cells in the granular layer of the
cerebellar cortex, which in turn extend their axons into the
molecular layer, where it divides and continues as the
parallel fibers. The Purkinje cells are the only output of
cerebellar cortex which project to the deep cerebellar
nuclei. Since the Purkinje cells are GABAergic, the output
of the cerebellar cortex is absolutely inhibitory. However,
the deep cerebellar nuclei receive excitatory inputs from
are mostly neuromodulators in function are also
said to set the mood of the individuals. Owing to
its reciprocal connections of RF to almost all brain
parts similar to the claustrum, it is believed to be
the “seat” of consciousness – from where the state
of consciousness is controlled and maintained.
Damage of RF creates coma-like conditions. RF
also has reciprocal connections to autonomic cen-
ters like hypothalamus and in lateral horn of spinal
segments, through which, it controls the ANS and
the endocrine system. RF also has important role
in generation and maintenance of sleep, rhythmic
control of respiration and cardiovascular func-
tions, and maintenance of tone of the axial and
11
the collaterals of the mossy and climbing fibers. The
Purkinje cell inhibition of the deep nuclei serves to modu-
late the level of this excitation. This inhibitory activity of
Purkinje cells is modulated by the interneurons like Basket
cell (on the body) and stellate cell (on the dendritic shaft)
from the molecular layer; both of the later cells receive
excitatory inputs from the parallel fibers. The molecular
layer contains the apical dendrites of a cell type called
Golgi cells; these neurons have their cell bodies in the
granular cell layer. The Golgi cells receive excitatory
inputs from the parallel fibers and in turn provide an
inhibitory feedback to the granule cells. The output from
the deep cerebellar nuclei reaches the thalamic nuclei and
then to the motor cortex modulating there ongoing neuro-
nal activity)
limb muscles. Sleep is identified with reduced
activity in RF neurons, in turn, less activation of
the cortex through RAS. Sleep also helps to refill
depleted store of aminergic neuromodulators. It
has a modulatory role in pain as ascending
spinothalamic fibers send collaterals to RF.
Numerous nuclei of RF create polysynaptic cir-
cuits for certain reflex behaviors like chewing and
eating, breathing, and head-neck movements in
response of a visual or auditory stimulus.
12
Autonomic Nervous System (ANS): Prime
Controller of Adaptive Behavior
Autonomic nervous system (ANS) is known to
have very important role in innate as well as
learned behavior. ANS has the sympathetic and
parasympathetic components and bears a larger
bodily spread than CNS. Hypothalamus presents
the highest center of the ANS with vaguely dis-
tinguishable sympathetic and parasympathetic
components. Brain stem contains the parasympa-
thetic outflow only. Spinal cord contains
thoracolumbar sympathetic outflow and sacral
segment parasympathetic outflow. ANS is
engaged in control of the animal behavior, by
resetting hypothalamic circadian clock in supra-
chiasmatic nuclei (SCN) and the neuroendocrine
balance along the HPA axis, in turn, influencing
neuroeffector communication at the peripheral
targets in response of the external or internal stim-
ulus. Hypothalamic autonomic nuclei send recip-
rocal connections to spinal autonomic centers.
Innate behaviors like hunger, thirst, and sexual
behavior are known to be regulated by contrasting
influences of sympathetic and parasympathetic
components of the ANS. Sympathetic component
of the ANS is said to be greatly involved in the
adaptive behaviors which present survival chal-
lenges like fight, flight, and fright situations
(Kumar et al. 2018c). An activation of sympa-
thetic component is associated with aggression,
anxiety, panic reaction, and disturbed appetite; in
contrast an activation of parasympathetic compo-
nent may be associated with ease of the mood,
relaxation, good appetite, sexual excitation, and
orgasm.
Emerging Behavioral Roles for Other Brain
Parts and Nervous System Components
Insula and “Mirror Neurons”: In Awareness of the
Self-Action and That of Others
Insular cortex which is anatomically buried
between frontal, temporal, and parietal cortices
is said to be involved in introspection and aware-
ness of the “self-actions” (Namkung et al. 2017).
A specific type of neurons, “mirror neurons,”
which are said to be present in anterior part of
the insula (and also in frontal, temporal, and
Neural Control of Behavior
parietal cortices surrounding it), and ACC are
believed to be principle to the awareness of the
“self-actions” (Ramachandran 1995). Mirror neu-
rons are also present in premotor regions of pre-
frontal cortex, temporoparietal junction (TPJ), or
inferior parietal lobule (IPL). These neurons were
first reported in macaque monkeys, when it was
found that these neurons activated not only during
performance of a task but also when a monkey
was made to observe others doing it. Mirror neu-
rons were also said to be behind social behaviors
like understanding what others think or feel
(hence in empathizing) (Carrillo et al. 2019), or
learning by imitation of others’ actions
(Ramachandran 1995).
Habenula: In Reward-Based Decisions and
Adaptive Behavior
Habenula is involved in the regulation of release
of neurotransmitters and modulators from brain
stem aminergic nuclei which is thought to be a
central mechanism for developing an adaptive
response to a persistent stimulus. Habenula is an
evolutionarily conserved bilateral structure
located at the interface of the diencephalon,
basal nuclei, and brain stem which works as a
gateway between cortical-subcortical and brain
stem structures (Kiernan et al. 2014; Kumar
et al. 2017b). It is thought to function as a switch-
board for regulating emotional behavior in condi-
tions facing survival challenges (e.g., to avoid or
approach or submission or freezing in response of
a stimulus). Medial and lateral components of the
habenular pathway are involved in positive and
negative reward prediction, respectively, making
their balanced action crucial for decision-making
in reward-oriented behavior (Kumar et al. 2017b).
Any adaptive behavior also involves modulation
in the settings of the hypothalamo-pituitary-
adrenal (HPA) axis.
Claustrum: In Unified Perception and
Consciousness
Claustrum is a thin sheet of gray matter
sandwiched between insular cortex and striatum
(it is separated from the insular cortex by the
extreme capsule and from the striatum by the
external capsule). Its unique location allows it to
Neural Control of Behavior
connect to all parts of the cerebral hemisphere
reciprocally. It also sends reciprocal connections
to contralateral hemisphere and other brain parts
as hippocampus, amygdala, basal nuclei, thala-
mus, brain stem, and nearly all other parts of the
brain. Uniquely, contralateral claustrums also
connect with each other through interclaustral
fibers. Empirical studies are scarce on the func-
tional role of claustrum. Any direct role in behav-
ior is not known, but recently proposed functions
to claustrum suggest that it creates a neural base
for production of any behavior. A study in the
epileptic patients which placed electrodes in at
one site of the white matter tract beneath the
claustrum made the patient to stare blankly
ahead with no recall of event when stimulus was
removed (Koubeissi et al. 2014). The above said
observation suggested mechanistic role of claus-
trum in the generation of consciousness. Claus-
trum fibers topographically connect with the
deeper layers of cortex (mostly layer six). Claus-
tral efferents to the cortex mostly end at the inter-
neurons hence indirectly bear inhibitory influence
on the pyramidal neurons. Inhibition of the cortex
by the claustrocortical fibers is said to have mod-
ulatory effect on the electrical activation pattern of
the cortical segments (Jackson et al. 2018). Stud-
ies also suggested its role in real-time synchroni-
zation of electric waves and oscillatory coupling
of two cortical regions bound to a common cog-
nitive function (Narikiyo et al. 2018). Francis
Crick, based on its universal cortical connections
(and also to the other brain parts) and role in
coordination of the cortical electrical activity, pro-
posed its role in generation of consciousness and
being conductor of the information flow through
the cerebrum (like conductor of an orchestra who
directs whole band!) (Crick and Koch 2005).
Function of claustrum resembles to an electrical
device which performs multimodal integration of
the stimulus information across the brain parts
hence helps to create a unified sense of perception
and generates a continuous flow of consciousness.
Enteric Nervous System (ENS): Mediator of the
Gut-Motivated Behavior
ENS-CNS connection has given rise to concept of
gut-brain axis. ENS is predominantly autonomic
13
in function, and in concert with the gut hormones,
it helps to maintain mood and emotion of the
individual. New research has established its role
in reward-mediated behavior. Uniquely,
enteroendocrine cells in the gut mucosa were
found to connect to the vagus nerve which is a
parasympathetic nerve supplying the gut (Han
et al. 2018). Right vagus nerve which innervates
gut was found to carry gut impulses to the dopa-
minergic nuclei in the brain stem – which present
important component of reward pathway
connecting to the forebrain (Han et al. 2018).
Parkinson’s disease, a neurodegenerative disease
of nigrostriatal pathway in the brain, is believed to
start in ENS neurons of the large gut. A
dysregulation of gut-brain axis has also been
found implicated in the psychological stress and
mood disorders like anxiety and depression.
Behavior-Specific Neural Correlates
Simple Behavior
Limb Reflex
A reflex is a rapid and involuntary response to a
stimulus or cue. It is controlled at the level of
spinal cord and is considered as a protective
mechanism to a suddenly presented threatening
stimulus (which is either painful and/or can poten-
tially injure the organism). A reflex is generated
by a sensory-motor circuit of spinal neurons
(Fig. 4) and doesn’t involve a cortical control.
Certain of the reflexes are also controlled at the
level of brain stem; these are head and neck move-
ments, mastication, and vital bodily functions
such as cardiovascular circulation and respiration.
Complex Behavior
Emotional Behavior
Emotional behavior is said to be generated and
controlled by the limbic system in the brain. Lim-
bic system comprises a rim of cortical mass sur-
rounding the diencephalon – also called as limbic
lobe, a set of subcortical nuclei – and white matter
connections between them (Fig. 5) (Kiernan et al.
2014). Amygdala, a quasi-cortical gray matter
14
Neural Control of Behavior, Fig. 4 A limb reflex arc
(Hammer tap stretches tendon which in turn stimulates
sensory receptors (spindles) in the agonist muscles. Sen-
sory neurons (blue) synapse with and excite motor neurons
(brown) in the ventral horn of the spinal cord. The motor
neurons innervate agonist muscle fibers (brown, intact
mass, is one of the main components of this sys-
tem. Amygdala is also called a center for fear,
aggression, anxiety, and panic, or more correctly,
it predicts salience value for any emotional infor-
mation (Janak and Tye 2015). A connection
between amygdala and prefrontal cortex is said
to exert control over impulsive behavior where
prefrontal control is inhibitory to amygdala, and
their relationship is antagonistic (Volman et al.
2011).
The neural circuitries related to memory, cen-
tered at hippocampus, are extensively connected
to the limbic system (as memory contents make
essential ingredients of any emotional behavior).
Also, autonomic and endocrine systems, which
are centered at hypothalamus, are intensively
linked with the limbic system, and their associated
activations contribute to the physical and psycho-
logical manifestations of any emotional behavior.
Neural Control of Behavior
line), and their stimulation causes immediate contraction
of these muscles. Sensory neurons also excite spinal inter-
neurons (purple) which in turn synapse to and inhibit motor
neurons innervating the antagonist muscles (brown, bro-
ken line), causing their relaxation)
Reward-Driven Behavior
Mesolimbic or reward pathway – a dopaminergic
circuitry – connects ventral tegmental area (VTA)
in the midbrain to ventral striatum or nucleus
accumbens (NAc) in the forebrain (Kiernan et al.
2014). This is the prime circuitry (VTA-NAc)
involved in all kinds of reward-based behavior
including natural responses for food and sex and
social interactions (Fig. 6).
This pathway is intensely connected to various
cortical and subcortical regions, hippocampus,
and brain stem nuclei. Prefrontal cortex keeps
this circuitry inhibited and hence puts a break on
reward-based behavior. This circuitry is involved
in the addictive behavior and is also implicated in
obsessive-compulsive disorders. Reward pathway
is evolutionarily conserved among animal species
which is based on the release of dopamine when
presented with a desirable stimulus. Same cir-
cuitry is involved in aversive behavior and stress
where a reduction in dopamine release brings
displeasure.
Neural Control of Behavior
Neural Control of Behavior, Fig. 5 Limbic system
(The limbic system is a complex set of structures found
on the central underside of the cerebrum, comprising inner
sections of the temporal lobes and the bottom of the frontal
lobe. It combines higher mental functions and primitive
Neural Control of Behavior, Fig. 6 Reward pathway
(The reward pathway is principally a dopaminergic path-
way (yellow, solid line) that connects ventral tegmental
area in midbrain to nucleus accumbens of ventral striatum
and the prefrontal cortex of cerebrum. Apart from dopami-
nergic efferents, the nucleus accumbens also receives
15
emotion into a single system often referred as the emo-
tional nervous system. The primary structures within the
limbic system include the amygdala, hippocampus, thala-
mus, hypothalamus, basal ganglia, and cingulate gyrus)
inputs from the Glutamatergic neurons (red, broken lines)
of the hippocampus, amygdala, and medial prefrontal cor-
tex. After being activated by glutamatergic inputs, GABA
release (black, broken lines) occurs onto the ventral
pallidum, consequently causing cortical disinhibition)
16
Reward system works on the prediction error
signaling (den Ouden et al. 2012). Reward cen-
ters, which contain dopamine-releasing nuclei,
generate prediction for the expected reward fol-
lowing behavior. A prediction error is computed
by these nuclei deducting the prediction estimate
from the actual reward value (den Ouden et al.
2012). When this actual reward is more than what
was expected, a positive prediction error is gener-
ated which causes release of dopamine – causing
pleasure and in turn enforcing/motivating the
individual to repeat the behavior. When actual
reward is less than expected, a decrease in dopa-
mine release (depression) is to occur – causing
displeasure and in turn aversion from that behav-
ior. When expectation is fulfilled, there has to be a
normal release, causing no/or little enforcement to
repeat the behavior. There is hierarchical differ-
ence in coding of reward along the reward path-
way. The neurons at the down most center,
SNc/VTA, activate with any novel information
irrespective of its nature (i.e., each novel informa-
tion is rewarding to the subject) based on its
saliency. Neurons in NAc encode both saliency
and valence of any incoming information. PFC
encodes saliency as well as valence of any incom-
ing information though its components exert dif-
ferential influence on the reward pathway.
Neurons in VMPFC/OFC activate more in posi-
tive prediction signaling, i.e., reinforce a reward-
ing behavior, and in DLPFC activate more in the
negative prediction signaling, i.e., create aversion
for a non-rewarding behavior (Fouragnan
et al. 2018).
Adaptive Behavior
Adaptive behaviors, like response of the individ-
ual to a threatening or rewarding stimulus or situ-
ation like success or failure, win or defeat, have a
complex neural circuitry which chiefly involves
medial and orbital PFC, ACC, and anterior por-
tion of the insula. Also are involved in this, corti-
cal and subcortical components of the reward
pathway, septal nuclei, and anterior hypothalamic
nuclei. Emerging evidence suggests master role of
the habenula in regulation of the adaptive behav-
ior (has been described in detail in section
Neural Control of Behavior
“Emerging Behavioral Role for Other Brain
Parts and Nervous System Components”).
Miscellaneous: Love, Lust, Hate, Disgust, Religious/
Ideological Fundamentalism, and Crowd Behavior
Though available knowledge on neural correlates
of these behaviors is still limited, functional neu-
roimaging studies put some light on the brain parts
grossly involved. Affectionate love is known to
activate specific cortical and subcortical compo-
nents of the limbic system which is overlapped but
effectively distinct from the sexual attraction. The
core brain regions that activate in a loving behav-
ior are medial PFC, ACC, anterior insula, head of
caudate nucleus or putamen, and VTA (Diamond
and Dickenson 2012). In sexual attractions, acti-
vation of caudate nucleus and VTA is less distinc-
tive, and an additional activation of the amygdala
and hypothalamus occurs (Diamond and
Dickenson 2012). Showing hateful and disgust
behaviors also causes activation of the amygdala.
Hateful behavior similar to the loving causes acti-
vation of the putamen and insula but, in contrast,
causes deactivation of frontal, temporal, and pari-
etal cortices (Zeki and Romaya 2008). Reward
(or aversion) pathway is activated both in loving
and hateful (or showing disgust) behavior. More
ventral regions of the striatum and pallidum get
involved in rewarding behavior like lust or aver-
sive behaviors like hate or disgust. Increase in
secretions of oxytocin by the hypothalamic nuclei
(Heinrichs et al. 2009) and neurotrophins by cor-
tical neurons is known during loving behavior
(Emanuele et al. 2006). Religious and ideological
fanaticisms (fundamentalisms) are known to
cause more activation of medial/ventral prefrontal
cortex with relative silencing of DLPFC. DLPFC
is thought to have role of a critic and activates
more in a rational behavior – when individual uses
reasoning to select a response (Harris et al. 2009).
Crowd behaviors involve complex interactions
between prefrontal cortex and limbic system com-
ponents, with a lesser prefrontal cortex-mediated
inhibition of the amygdala (Huis in ‘t and de
Gelder 2015). An added influence of oxytocin
and vasopressin (secreted by supraoptic and para-
ventricular nuclei of the hypothalamus, respec-
tively) on decision-making by individual
Neural Control of Behavior
members has been evidenced in such behaviors
(Heinrichs et al. 2009).
Neural Basis of the Pathological Behaviors
Brain parts involve differentially in various
known pathological behaviors. A dysregulation
of reward pathway is associated with many path-
ological behaviors like addiction, substance
abuse, and gambling (Yun et al. 2016). An
amygdala- prefrontal imbalance has been associ-
ated with aggression, anxiety, depression, and
panic reaction and the diseases like major depres-
sive disorder (MDD) and bipolar disorder (BPD)
(Volman et al. 2011). MDD and BPD have also
been linked with altered activity of hippocampal
neurons. An abnormal activity at temporoparietal
cortical junction (TPJ) has been associated with
hallucinations in schizophrenia. A hyperactivity
of the dopaminergic neurons in reward pathway
has been noted in psychotic disorders.
A dysregulated brain plasticity also has been
attributed for the pathological behavior, like
excess anxiety, aggression, panic, or post-
traumatic stress disorder (PTSD) (Kays et al.
2012). Social defeat and self-harming behaviors
involve experience induced dysregulated plastic-
ity in the brain regions involved in adaptive
behavior like prefrontal cortex, amygdala, and
aminergic nuclei in the brain stem. Dysregulation
of the basal nuclei circuitry has been suggested in
obsessive-compulsive disorders (OCDs).
Dysregulation of the neural circuitry involved
in adaptive behavior also can give rise to certain
behavioral pathologies. Repeated social defeat
may induce a state of chronic unpredictable stress
(CUS) and learned helplessness (a behavioral
state in which individual no more explores for a
safer option) in the individuals. CUS may be a
reason for self-harming behavior seen in some
individuals – which supposedly give them some
sense of release from a conflicting condition or
mental strain (Blasco-Fontecilla et al. 2016).
Owing to the increasing daily life use of new
information technology facilitated digital devices
like computers and smartphones, certain new
behavioral disorders have emerged. “Problematic
use of the Internet (PUI)” is an umbrella term for
such disorders which involves “Internet video
17
gaming and pornography,” excessive social
media network use, and selfitis – an excessive
preference for selfie taking and posting it on social
media. Though many of the PUI-affiliated disor-
ders are still not recognized by Diagnostic Statis-
tical Manual (DSM) of psychiatry, emerging
research suggests these behaviors involve reward
system of the brain quite similar to other recog-
nized addictive behaviors (Kumar et al. 2017a).
Cross-References
▶ Behavior Systems
▶ Endocrine System
▶ Heritability of Behavior
▶ Hormones and Behavior
▶ Model Fitting
▶ Theory of Mind
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