MAJOR REVIEW A C TA Vol. 18, No.

1, 2020, 1-14
NEUROPSYCHOLOGICA
Received: 11.10.2019
Accepted: 26.02.2020 CURRENT VIEWS
A – Study Design ON NEUROPLASTICITY: WHAT
B – Data Collection
C – Statistical Analysis
D – Data Interpretation IS NEW AND WHAT IS OLD?
E – Manuscript Preparation
F – Literature Search
G – Funds Collection
Bożydar L.J. Kaczmarek
University of Economics and Innovation, Lublin, Poland

SUMMARY
The main aim of the paper is to show that many previously
forgotten discoveries within the field of neuroscience own
their rediscovery and renaissance to the refinement of tools
provided by the technological advances. Most spectacular is
the advancement of brain imaging techniques, which provide
hard data that support for evidence for previously neglected
presumptions and ideas. Neuroplasticity is an example of
such a long ignored historical discovery. One reason for that
neglect is that it stood in contradiction to beliefs and theories
prevailing at the first half of the twenties century. The idea
of neuronal plasticity is not disputed any longer since it has
found confirmation not only in a dramatic development of
neuroimaging but also in the advancement of neurobiology.
Most authors concentrate upon neuronal plasticity, recent
studies, however, have produced a wealth of information re-
garding neurogenesis, in which astrocytes have proved to
play a significant role. The significance of adult neurogenesis
for learning and memory and for treatment of depression is
outlined. Moreover, it was observed that neuroplasticity ben-
efits patients suffering from obsessive-compulsive disorder
(OCD) who undergo effective, evidence-based treatment.
Convincing examples of brain plasticity brings also clinical
practice, which often unveils the appearance of hitherto hid-
den artistic abilities in people who have suffered from brain
damage. In addition, the possibilities of altering the brain
functions by mental force alone are discussed. Thus, the
paper reveals that many “controversial” ideas were con-
firmed by contemporary studies forcing changes in a tradi-
tional view on brain works.

Key words: research paradigms, neuroplasticity,

glial plasticity, mental force, neurogenesis

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 Kaczmarek, Current Views on Neuroplasticity

The present paper concentrates on previously forgotten discoveries within

the field of neuroscience. It is stressed that their rediscovery and renaissance
have been due to the technological advances of brain imaging techniques over
the past decades. Most spectacular is the advancement of brain imaging tech-
niques, which provide hard data that support evidence for previously neglected
presumptions and ideas. Neuroplasticity is an example of such a long-ignored
historical discovery. One reason for that neglect is that it stood in contradiction
to theories and research paradigms prevailing in the first half of the twenties cen-
tury. It is worth to remind that research paradigms are built upon our mental
schemas or models of objects and events that surround us. Since our senses
are pervaded by an abundance of stimuli the brain selects only some of them to
create a coherent and comprehensible image of the reality. The world’s image
consists of mental models, which in fact are simplified schemas encompassing
the most important features of a given object, event or action. Contrary to a gen-
eral belief, we do not perceive ready-made information. Indeed, our perception
is an act of creation and not a synthesis of particular features. Put differently, the
images or models (percepts) are created in a process that strongly resembles
sculpting: delineating distinctive features of an object or event (Brown, 2010,
2015; Dehaene, 2013; Pąchalska, Kaczmarek, and Kropotov, 2014). Those mo-
dels play an important role both in perception and in action. Moreover, they pro-
vide stability and make the world safe and comprehensible.
Research paradigms are specific types of mental models, and, consequently,
they provoke the scientific community to reject ideas that stand in contradiction to
existing theories. Kuhn (1996) argues that research paradigms provide a stiff intel-
lectual framework, or a “disciplinary matrix”, imposing the ways of data gathering
and interpreting the results. In consequence, any discrepancies are considered to
follow from experimental errors or anomalies. Over time more and more anomalies
and unsolved questions accumulate, and it turns difficult to solve them within
a currently existing paradigm. As a consequence a paradigm shift takes place
and a new model of thought aimed at explaining the existing puzzles is created.
An example of such a shift is an approach to the idea of neuroplasticity.

Neuronal Plasticity
Neuroplasticity is extensively discussed in the contemporary literature, and it
is often viewed as a recent discovery. Therefore, it might be worthy to remind
experiments performed already at the beginning of the twentieth century. Two
prominent physiologists, Graham Brown and Charles Sherrington, stimulated the
motor cortex of experimental animals and found that the stimulation of the iden-
tical place in a particular monkey produced twitches of different muscles. Skep-
tics argued, however, that differences among movement maps of animals might
well be inborn and not acquired (Schwartz, and Begley, 2003). This assumption
was dispelled by Karl Lashley in 1923, who derived different maps in the brain
motor area of one particular rhesus monkey. Lashley was able to show that those
maps underwent considerable changes over the course of one month, in the

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wake of different motor tasks the monkey had performed. The idea was further
supported by Donald Hebb (1949), who argued that learning and memory depend
upon the strengthening of the activity of synapses in order for two events or stim-
uli to be bound together.
Those findings were neglected due to the prevailing belief in the immutability
of the central nervous system. Researchers promoting a possibility of changes
in neuronal circuits were as a rule attacked by the scientific establishment. It suf-
fices to mention Walther Spielmeyer and Max Bielchowsky two widely recognized
German neuropathologists (Stahnisch, and Nitsch, 2012). Spielmeyer collabo-
rated with renowned scholars as Alois Alzheimer, Korbinian Broadmann, Emil
Kraepelin, and Franz Nissl, while Bielschowsky is best known as the inventor of
a derivative silver staining technique (Stahnisch, 2015). This might explain the
ambiguous position of Santiago Ramón y Cajal, the great Spanish anatomist
and Nobel laureate. In his Nobel lecture delivered in 1906, he firmly stated that the
pathways in he firmly stated that the pathways in the mature brain are fixed and
immutable (Kolb, 1995). In his later works, however, he assumed a possibility to
elicit new connections between neurons in the process of learning (Stahnisch,
and Nitsch, 2012; Pąchalska et al., 2014).
The revival of the notion of neuroplasticity took place thanks to experiments
of Merzenich and his co-workers (1983) on changes within cortical somatosen-
sory fields. As is often the case, Merzenich’s initial observations met with strong
opposition. It was argued that the expansion of cortical areas of the hairy back
of the hand, which followed cutting the nerves of the palm, might be a result of
unmasking already existing connections. To convince their critics, Merzenich at
al. (1984) decided to take a drastic step amputating an owl monkey’s finger to
cut off all sensory input to it. After a couple of months, they then recorded the elec-
trical activity of the monkey’s somatosensory area. The recording showed that cor-
tical representation of the amputated finger was taken over by the adjacent digits
of the palm. It confirmed the possibility of changing the cortical representation of
fingers of the hand. In 1991 Allard, Jenkins, and Merzenich created artificial syn-
dactyly by sewing together two adjacent digits of adult owl monkeys. After two to
three months they compared the previously recorded somatosensory brain maps
of the monkey’s hands with the recordings following the surgery and found that
the two sewn digits had now a single overlapping representation.
At the same time, Taub (1980) showed that the deafferentation of one of a mon-
key’s arm resulted in an inability to perform voluntary skillful movements using the
limb. He then immobilized the monkey’s healthy hand forcing it to use the injured
one. Such training resulted in a significant improvement in the efficiency of in-
jured hand movements. It is notable that Taub drew on observations published
by German physiologist Hermann Munk already in 1909 (see Doidge, 2007).

Clinical Significance of Brain Plasticity

In humans, neuroplasticity also benefits patients suffering from obsessive-
compulsive disorder (OCD) who undergo effective, evidence-based treatment.

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 Kaczmarek, Current Views on Neuroplasticity

Schwartz and Begley (2003) reported metabolism alternation in the OCD brain
circuit after cognitive-behavior therapy had been applied. Positron emission to-
mography (PET) revealed a decrease of abnormally high hyperactivity of brain
structures of that network (mainly the caudate, the orbital frontal cortex, and the
cingulate gyrus).
Yet – as we all know – the cognitive-behavior therapy requires a long, effortful,
and distressing work of both the patient and the therapist. Besides, the results
are not always satisfactory. It raises one important question. Why is it so difficult
to change our habits if the brain is believed to undergo constant changes? It
needs to be noted that plasticity does not mean instability. Indeed, our everyday
actions require stability to be performed in an efficient way without having to think
about each individual movement. Repeating a particular action brings about the
strengthening of connections among groups of neurons responsible for the ac-
tion’s performance. In consequence, those inter-neuronal connections become
stable just like ruts in an old road. It means that repetition leads to rigidity by
forming some kind of mental (and neuronal) “tracks”, from which are difficult to
extricate (Doidge, 2007; Schwartz and Begley, 2003).
Nonetheless, improvement of brain-damaged patients who were given care-
fully planned treatment including art therapy confirms the possibility of overcom-
ing those difficulties. Moreover, the hitherto hidden artistic abilities are quite often
revealed in the course of treatment (Pąchalska et al., 2014). It is worth pointing
out that some of the patients reached a high level of their art of painting as was
the case in a patient diagnosed with schizophrenia (Pąchalska et al., 2008). He
was run over by a car after rushing out into traffic while actively hallucinating that
he could fly. In effect, he suffered a brain contusion and remained unconscious
for about 5 hours. After recovery he presented with disturbances of working
memory, hemispatial neglect, and perseveration (Pachalska 2007; 2020). The
tendency to perseverate was reflected in a series of his early paintings, which
were composed of three overlapping human figures (see Fig. 1).
He has since become a prolific artist, who exhibits his paintings all over the
world. At the same time, his works receive very good opinions from art critics
(see Pąchalska et al., 2013, 2014). The critics also point out that his recent paint-
ings progress towards symbolic art as exemplified in Figure 2.
The above-mentioned case, as well as the artistic achievements of other
brain-damaged patients, seem to support the idea of neural Darwinism proposed
by Gerald Edelman (1987). He presumes that the brain neuronal networks com-
pete with each other to perform a given task. In consequence, the job is done by
the network that performs the work best. Hence, any function may be taken over
by other brain networks if the one (or some) of them has been impaired. That
idea finds additional support in an experiment conducted by Pascual-Leone and
co-workers (2000). The authors disabled the occipital lobes of blind persons with
electromagnetic coil (Transcranial Magnetic Stimulation), and found that they
lost fluency in reading Braille texts. It shows that in the blind visual occipital cortex
plays a significant role in reading the Braille alphabet.

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 Kaczmarek, Current Views on Neuroplasticity

Fig. 1. Perseverated paintings produced by a patient diagnosed with schizophrenia

Source: Pachalska, Kaczmarek, Kropotov (2014)

Fig. 2. A recent symbolic painting of the same painter

Source: clinical material of M. Pachalska

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 Kaczmarek, Current Views on Neuroplasticity

The Power of Mental Force

Pascual-Leone et al. (1995) performed also an experiment that displayed the
possibility of reconstructing neuronal circuits with the use of mental exercise
alone. They trained two groups of people in skills of playing the piano. The first
group’s participants were asked to imagine they played and listened to a one-
handed, five-finger piano exercise for two hours, for five days. The other group
was actually playing the same sequence during the same period. Changes in
cortical areas targeting fingers involved in the task were observed in both groups.
Furthermore, the changes were limited to the cortical representation of the hand
used in the exercise. The researchers noted the improvement of playing skills in
both groups, although, naturally, the improvement produced by physical exercise
was better. Yet, a two hour physical exercise resulted in a considerable improve-
ment of the playing skills of the mental practice group.
Some neuroimaging studies have shown that mental and motor training acti-
vate distinct neural networks (Graybiel et. al., 1994; Lacourse et al., 2005; Nyberg
et al. 2006; Olsson et al., 2008), while others indicate that the two types of train-
ing can affect similar, but not identical, neural circuits (Decety, 1996; Decety et. al,
1989; Jeannerod, 1994, 1995; Kosslyn et al., 2007; Lotze et al., 1999; Munzert et
al., 2009). Slagter et al. (2011) argue that motor imagery depends on cognitive
higher-order aspects of action control, and report studies that found higher activa-
tion of midline and lateral motor as well as midline frontal areas (see also Nyberg
et al., 2006; Ranganathan et al., 2004). This research attests to the possibility of
direct influence on motor networks, which might explain the improvement of per-
formance following mental exercise. It might be of interest to remind the reader
that such a possibility was presumed by Raymon y Cajal over one hundred years
ago (see Doidge, 2007).
Recent years brought additional examples of changes in the brain work in-
duced by meditation practice. Neuroimaging studies using functional magnetic
resonance (fMRI) and positron emission tomography (PET) scans have shown
that the changes include both alterations in connectivity among the various brain
networks as well as structural changes (Brefczynski-Lewis et al., 2007; Davidson
and Lutz, 2008; Fan, 2011; Lazar et. al., 2005; Lutz et al. 2008, 2009; Slaghter
et al., 2011; Telles et al. 2015).
Fox et al. (2014) performed a meta-analysis on 21 brain imaging studies ex-
amining 300 meditation practitioners. They found:

…eight brain regions consistently altered in meditators, including areas key

to meta-awareness (frontopolar cortex/BA 10), exteroceptive and interocep-
tive body awareness (sensory cortices and insula), memory consolidation
and reconsolidation (hippocampus), self and emotion regulation (anterior and
mid cingulate; orbitofrontal cortex), and intra- and interhemispheric commu-
nication (superior longitudinal fasciculus; corpus callosum). (p. 40)

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 Kaczmarek, Current Views on Neuroplasticity

A follow up by Fox et al. (2016) suggests that different meditation styles are
reliably associated with distinct brain activity. Activations in some brain regions are
usually accompanied by deactivation in others but convergence is the exception
rather than the rule. They conclude that the results should be interpreted with cau-
tion since theoretical bias as well as methodological limitations are of concern.
The above described studies concentrated on neuronal circuits in accordance
with the “neuron doctrine” supported by Ramón y Cajal in his disputes with Golgi
(Koob, 2009; Stahnisch and Nitsch, 2012). Recent research, however, shows
that it is not only restructuring the neuronal networks that plays a significant role
in brain plasticity but also so far neglected glia.

Glial Plasticity
Preoccupation with neurons had its roots in discoveries of electric capacities
of the human body and hence the supreme importance of electrical signaling in
neurons. In the eighteenth century, the interest in electricity was comparable to
the fascination with quantum physics today. The interest was beefed up by elec-
tro-stimulations provoking contraction of muscles in dead bodies. Those demon-
strations, often applied in a highly theatrical way, were meant to show the power
of galvanism. Suffice it to mention the experiments of Giovanni Aldini (see Parent,
2004) who triggered contractions of various muscles by applying currents at dif-
ferent points along the head and body. Remarkably, he was able to produce ex-
pressions of smiling or grimacing in this way.
Glial cells were found to be electrically nonexcitable; hence they were ex-
cluded from further research. Recent studies, however, revealed that neurons
do not function in isolation. In fact, they are embedded in complex glial networks
in which astrocytes appear to play a very important role. They modulate multiple
aspects of synaptic plasticity by releasing gliotransmitters, calcium signaling, and
oscillations, coupling via gap junctions. Moreover, glia stabilize new synapses and
provide ensheathment of synapses, which restricts the spread of neurotransmitter
to neighboring synapses (Barker and Ullian, 2010; Chao et al., 2002; De Pita et al.,
2016; Kandel et. al., 2016; Pirttimaki and Parri, 2013). In addition, astrocytes maintain
hemodynamic control, metabolic demand homeostasis, and energy supply (De Pitta,
2016; Giaume et al., 2010; He and Sun, 2007; Navarrete and Araque, 2011).
Above all, astrocytes have proven to be able to communicate among themselves
via calcium waves, which results in a release of glutamate, and is linked to
hemodynamic control. They form complex communication nets that can generate
various regulatory signals and bridge structures and networks that are otherwise
disconnected from each other (Fields, 2010; Volterra and Meldolesi, 2005).
Barker and Ullian (2010) state that:

Astrocytes are organized into discreet domains, with the territory of a single
astrocyte estimated to contact between 300 to 600 dendrites and upwards
of 105 synapses…This extensive synaptic interaction …positions astrocytes
to directly influence the structure and function of the synapse. Indeed, there

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 Kaczmarek, Current Views on Neuroplasticity

is increasing evidence that astrocytes play active roles in controlling the

number and strength of a neuron’s synapses and thus may contribute to
mechanisms underlying synaptic plasticity. (p. 40)

It was the introduction of fluorescent calcium indicators that enabled revealing

the communicative role of calcium (Fields, 2010). Recent studies unveiled also
the role of sodium in neuron-glia interaction and neuro-metabolic coupling. This
is an important finding since cellular sodium homeostasis is of the upmost func-
tional significance for the brain. Indeed, most of the brain energy is consumed
by the Na+/K+-ATPases (Carter, and Bean, 2009; Howarth et al., 2012). Accord-
ingly, altered sodium homeostasis is observed in brain pathological conditions
(Rose, and Chatton, 2016).

Neurogenesis
We also cannot pass over neurogenesis, in which astrocytes have proved to
play a significant role. The neurogenesis (the ability to generate new brain cells)
takes place in the subventricular zone (SVZ), and the subgranular zone (SGZ) (Alt-
man, 1969; Doetsch at al., 1999; Luskin, 1993) as well as in hippocampus (Seneki,
et. al., 2001, Wang, and Bordey, 2008; Woollett, and Maguire, 2011). Generation
of new neurons is largely restricted to the adult SVZ and SGZ, whereas astro-
cytes and oligodendrocytes are continuously born throughout life (Duan et al.,
2008, Bruel-Jungerman, Rampon, and Laroche, 2007). Those new cells show
unique electrophysiological characteristics, which makes them prone to undergo
plasticity. It is a multi-step process modulated to a considerable degree by ge-
netic, behavioral and environmental factors. A significant role of those newborn
cells is facilitation of learning and memory due to the ability to incorporate into
existing neural circuits. Indeed, Woollett, and Maguire (2011) observed an in-
crease in the hippocampus volume in London taxi drivers. Significantly, no such
changes were observed in bus drivers who used to drive a constrained set of
routes (Maguire, Woollett, and Spiers, 2006)
On the other hand, environmental factors, stress in particular, result in a shrink-
age of dendrites or even cell death (Sapolsky, 2004). Stress hormones deplete
hippocampal neurons of glucose makings them sensitive to elevations of glutamate
to which they have a toxic reaction. Moreover, stress hormones have been found
to amplify the amygdala reaction to fear leading to a significant rise of hormones
and neurotransmitters such as cortisol, noradrenaline, and epinephrine. It mobilizes
body resources to deal with short-term dangers. In case of a prolonged stress,
however, the hypothalamic-pituitary-adrenal axis (HPA axis) is pathologically ac-
tivated leading to erosion of resilience and depletion of metabolic reserve (Le
Doux, 2002, Sapolsky, 2004). The malfunction of HPA axis is believed to be linked
to the development of depressive disorders (for review see Davey, Breakspear,
Pujol, and Harrison, 2017; Gerhard, Wohleb, and Duman., 2016).
Accordingly, treatment of depression is directed to alter the abnormal level of
cortisol. The therapeutic onset of antidepressant medication is delayed since the

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 Kaczmarek, Current Views on Neuroplasticity

drugs used here activate the second messenger systems. It leads to increased
gene activation and to a synthesis of proteins that enhance synaptic transmis-
sion. In effect, it enables new memory formations, learning new ways of coping
with stressful situations, and enhancing future learning experiences (Duman, et al.
2001, 2004). In other words, the greater plasticity of the brain is acquired. Since
such treatment takes time the research aiming at developing faster-acting thera-
peutic agents such as ketamine is being conducted (Duric, and Duman, 2013;
Gerhard, Wohleb, and Duman, 2016; Medrihan, et. al., 2017). Yet the outcome
of such procedures is still not sufficiently documented.

CONCLUSIONS
We remind the reader that new ideas do not suddenly appear out of a void in
the mind of the individual. In the case of science, it is the refinement of research
methods as well as an accumulation of the new data that make possible a new
interpretation of the existing but so far neglected theories (see also Kaczmarek
and Markiewicz, 2018). It also allows seeing the existing ideas in a new light as,
for example, the possibility of changing the brain functions by mental effort alone.
In addition, the emergence of new techniques created possibilities to determine
the contribution of astrocytes to brain plasticity, and their role as integrators of
neuronal network activity. Other significant manifestations of plasticity are alter-
nations in the brain works induced by meditation practice. Neuroimaging studies
using functional magnetic resonance (fMRI) and positron emission tomography
(PET) scans have shown that the changes include both alterations in connectivity
among the various brain networks as well as structural changes.
We also argue that brain plasticity is the rule and not exception. It enables
humans as well as animals to deal effectively with the changes occurring in the
environment, and, therefore, plays a significant role in learning. It can be best
observed in the hippocampus where new cells are born in response to new ex-
periences and training. Moreover, experimental evidence reveals a link between
synaptic plasticity and the proliferation and survival of new hippocampal neurons.
At the same time, sensitivity to environmental factors makes hippocampal cells
prone to react adversely to harmful situational agents such as heavy trauma or
stress leading to the hippocampus shrinkage. Hence, decreases in hippocampal
volume can be also observed in people plagued with depression. Spectacular
development of neurobiology made it possible to create new medications that
are successfully used in the treatment of depression. Those antidepressants
stimulate genes to create new proteins, and, consequently, cause alternations
in the level hormones and neurotransmitters.
In sum, the data presented in the paper prompt to revise the idea of neuro-
plasticity. It is linked not only with alternations within neuronal circuits, and neu-
rogenesis but with epigenetic changes as well. In fact, presently carried research
has marked a new era not only in neuroscience but also in genetics. Current epi-
genetic research has thrown a new light upon factors that affect gene activity

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and expression. In effect, the contemporary idea of neuroplasticity includes not

only cellular mechanisms but also environmental, and genetic factors. It also
means that our brain is prewired and not hard-wired to a considerable extent
(see Marcus, 2004; Merzenich, 2013).

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Corresponding author:
Bożydar L.J. Kaczmarek,
University of Economics and Innovation in Lublin (WSEI)
Projektowa 4
20-209 Lublin, Poland
e-mail: [email protected]

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