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Evolution of movement process as a key for human cognition.

  • Learning & Neurodevelopment Research Center


Movement is defined as a complex event in both the evolution of species and human development, which involves genetic and epigenetic mechanisms. It is linked with memory, attentional and linguistic processes, and it is required to create and use tools, defined as an extension and externalization of human hands, or the motor organs or effectors, so we believe is the basis of human cognition. The process created a parietal plasticity when incorporating tools into the body schema, which gave place to brain expansion by tooluse training. This sequence is considered relevant to the Homo sapiens development, and produces such level of sophistication to every cultural expression, that makes movement an important process both phylogenetic and ontogenetically. Under this context, this article covers the evolution of movement as a process. It begins with the first molecular actions to create a mechanism to retain energy and metabolize food. Additionally, this article explains: 1) how motility opened a door to the evolution of species, 2) how actin gets an important role in the cytoskeletal support, and 3) the development of the skills that allowed them to survive. Lastly, we investigate the evolution of movement as an adaptation to the environment, and the design of a human brain capable of pushing not only every muscle to the limit, but becoming part of other systems as memory, language or attention, as part of the cognitive processes on humans.
Evolução do processo de movimento como chave
para a cognição humana.
Evolution of movement process as a key for
human cognition.
Evolución del movimiento como clave para la
cognición humana.
Learning & Neuro-Development Research Center, USA. Alma Dzib-Goodin ORCID: 0000-0001-9705-9153. Daniel Yelizarov ORCID:
Correspondencia: Alma Dzib-Goodin. Learning & Neuro-Development Research Center, USA. Address: 6450 Cape Cod Ct Lisle, Illinois
60532. USA Email:
Alma Dzib-Goodin
Daniel Yelizarov
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
Cuadernos de Neuropsicología
Panamerican Journal of Neuropsychology
Movement is dened as a complex event in both the
evolution of species and human development, which
involves genetic and epigenetic mechanisms. It is linked
with memory, attentional and linguistic processes, and
it is required to create and use tools, dened as an
extension and externalization of human hands, or the
motor organs or effectors, so we believe is the basis
of human cognition. The process created a parietal
plasticity when incorporating tools into the body
schema, which gave place to brain expansion by tool-
use training. This sequence is considered relevant to the
Homo sapiens development, and produces such level of
sophistication to every cultural expression, that makes
movement an important process both phylogenetic and
Under this context, this article covers the evolution
Keywords: Movement; evolution; actin proteins; cell evolution;
Central Nervous System development; cognition; Memory;
El movimiento, desde la perspectiva evolutiva, es una
necesidad de las especies para sobrevivir sobre la faz
de la tierra, que involucra mecanismos tanto genéticos
como epigenéticos, vinculados a los procesos de
memoria, atención y lenguaje, necesarios para crear y
usar herramientas, empleadas como extensión de las
manos humanas y de los órganos motores o efectores.
Esto creó una plasticidad parietal al incorporar
herramientas en el esquema del cuerpo, y dio lugar a
la expansión del cerebro mediante el uso de las mismas
y el aprendizaje de cómo usarlas. Esta secuencia se
considera relevante para el desarrollo del Homo sapiens,
y convierte al movimiento en un proceso logenética y
ontogenéticamente importante.
En este contexto, este artículo cubre la evolución
del movimiento como proceso. Comenzando con las
Palabras clave: Movimiento; Evolución; Proteína actina;
Evolucn celular; Desarrollo del Sistema nervioso central;
Cognición; Memoria; Lenguaje.
O movimento, desde a perspectiva evolutiva, é uma
necessidade das espécies para sobreviver sobre a face
da terra, que envolve mecanismos tanto genéticos como
epigenéticos, vinculados aos processos de memória,
atenção e linguagem, necessários para criar e usar
ferramentas, empregadas como extensão das mãos
humanas e dos órgãos motores ou efetores. Isto criou
uma plasticidade parietal ao incorporar ferramentas
no esquema do corpo, e deu lugar à expansão do
cérebro mediante o uso das mesmas e a aprendizagem
de como usá-las. Esta sequência se considera
relevante para o desenvolvimento do Homo sapiens, e
converte o movimento em um processo logenético e
ontogenéticamente importante.
Neste contexto, este artigo coloca a evolução do
movimento como processo. Começando com as
Palavras-chave: Movimento; Evolução; Proteína actina;
Evolução celular; Desenvolvimento do Sistema nervoso
central; Cognição ; Memória; Linguagem.
of movement as a process. It begins with the rst molecular
actions to create a mechanism to retain energy and metabolize
food. Additionally, this article explains: 1) how motility opened a
door to the evolution of species, 2) how actin gets an important
role in the cytoskeletal support, and 3) the development of the
skills that allowed them to survive. Lastly, we investigate the
evolution of movement as an adaptation to the environment,
and the design of a human brain capable of pushing not only
every muscle to the limit, but becoming part of other systems
as memory, language or attention, as part of the cognitive
processes on humans.
primeras acciones moleculares para crear un mecanismo
capaz de retener la energía y metabolizar los alimentos.
Además, este artículo explica: 1) cómo la motilidad abrió una
puerta a la evolución de las especies, 2) el papel de la actina
en el apoyo al cito esqueleto, y 3) el desarrollo de habilidades
que permitieron la pervivencia de las especies. Por último,
se investiga la evolución del movimiento como adaptación al
medio ambiente, y el diseño de un cerebro humano capaz de
empujar no sólo cada músculo al límite, sino convertirlo en parte
de otros sistemas como la memoria, el lenguaje o la atención,
como parte del proceso cognitivo en los seres humanos.
primeiras ões moleculares para criar um mecanismo capaz
de reter a energia e metabolizar os alimentos. Ademais, este
artigo explica: 1) como a mobilidade abriu uma porta à evolução
das espécies, 2) o papel da actina no apoio ao cito esqueleto,
y 3) o desenvolvimento de habilidades que permitiram a
sobrevivência das espécies. Por último, se investiga a evolução
do movimento como adaptação ao meio ambiente, e o desenho
de um cérebro humano capaz de empurrar nâo só cada músculo
ao limite, mas o converter em parte de outros sistemas como a
memória, a linguagem ou a atenção, como parte do processo
cognitivo nos seres humanos.
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
Cuadernos de Neuropsicología
Panamerican Journal of Neuropsychology
We want to thanks especially to Bill Goodin, for his
amazing support, readings and correction of this paper.
Nature is ever at work building and pulling down, creating and destroying, keeping everything whirling and owing,
allowing no rest but in rhythmical motion, chasing everything in endless song out of one beautiful form into another”
John Muir
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
Cuadernos de Neuropsicología
Panamerican Journal of Neuropsychology
Movement process is a complex evolutionary feature
that began long time ago even before eukaryote cells.
Currently, cell movement is possible through different
process, which can be a somatic, biochemical, diffusible
or non-diffusible sign that can be detected by receptor
proteins located on the cell membrane and spread by them
through signaling forces to the cell (Ananthakrishnan and
Ehrlicher, 2007).
This process is possible thanks to a precise and
complicated network of genes, proteins, and enzymes
while engages a boundless redisposition of the actin
cytoskeleton, through three stages in most of cells. First,
a cell pushes the membrane forward by standing and
regrouping (growing) the actin network at its leading edge.
Second, it follows to the substrate at the leading edge and
deadheres (discharges) at the cell body and posterior
areas. Third, contractile energy, produced mainly by the
action of the acto-myosin network, pulls the cell forward
(Ananthakrishnan and Ehrlicher, 2007).
However, this process didn’t begin working this way
since the beginning of the nature evolution. At some early
point of life on this planet, it was required the introduction
of ATP as the universal energy, which was an important
stage in bioenergetic improvement. ATP synthase is
an enzyme that creates the energy storage molecule
adenosine triphosphate. ATP is the most commonly
used energy exchange of cells for all organisms. This
act displacing acetyl phosphate, which is a compound
related in taurine, pyruvate and hypo taurine metabolism
(Sousa, Thiergart, Landan, Nelson-Sathi, Pereira, Allen,
Lane, and Martin, 2013). However, even if ATP is found
through ancestries, it is not the only one motor inside of
individual cells. The most accepted justication for ATP’s
increase to become so important, it is because is a result
of the substrate specicity of the rotor stator-type ATPase.
This protein, is universal between cells as the sequencer
(Thauer, Kaster, Seedorf, Buckel & Hedderich, 2008) of
all biological energy in the form of ATP, and it is produced
from chemiosmotic pattern, so it has as work to protect
the separation from the inside of the cell to the outside,
and the harnessing of that electrochemical gradient via
a coupling factor, as an ATPase of the rotor–stator-type,
meaning it has a better chance to succeed and it was
adapted by most of cells (Martínez-Cano, Reyes-Prieto,
Martínez-Romero, Partida-Martínez, Latorre, Moya, and
Delaye, 2015).
However, these machineries were recruited long
before the modern eukaryote cell, because prokaryotes,
the rst alive organisms, were developed in a sheltered
and chemically rich medium, with dissimilar ways to get
energy in order to move. In this sense, protein kinase cyclic
nucleotide-binding (CNB) domains were widespread in
the prokaryotic world, so it is believed that they were an
earliest draft that co-evolved beside the cAMP (adenylyl
cyclase pathway) or, as a mechanism for translating the
stress-induced cAMP as a second messenger into a
biological reaction. There have been found some kinase
domains in prokaryote cell, so it is accepted that both
cAMP and cGMP domains could be functionally related in
the evolution of eukaryotes to an EPK (Eukaryotic Protein
Kinase), so they can be found, for example in all fungi
(Taylor, Keshwani, Steichen, and Kornev, 2012).
Assuming that motility is a process required to get
an integration of nuclear and other cellular functions as
a bidirectional passage across the nuclear envelope,
this of course requires that all tRNA, rRNA and mRNAs
must be re-distributed, and since proteins required for
DNA replication, transcription, transcriptional regulation,
RNA processing and overall nuclear organization are only
imported, since translation is cytoplasmic, (Wickstead
and Gull, 2011; Koumandou, Wickstead, Ginger, van
der Glezen, Dacks and Field, 2013; Blombach, Smollet,
Grohmann, Werner, 2016).
Under this idea, it is necessary to understand how
cells changed.
Motility: from Prokaryotes to Eukaryotes
There is not a consensus about how the rst cells
were originated, some data suggest that the eukaryotic
cell could appear from a merger of two prokaryotes cells,
but most compelling evidence specically mention an
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
Cuadernos de Neuropsicología
Panamerican Journal of Neuropsychology
archaeal host and a bacterial endosymbiont process that
could produce a new kind of nature item and may have led
to the contemporary complex eukaryotic cell (Davidov &
Jurkevitch, 2009).
Of course, that had to produce a number of new
mechanisms to get energy, so it is possible to say that
primitive eukaryote possible could become a predator
with the ability to devour bacteria and archaea in order
to get food, but eventually endosymbiosis would lead
to the improvement of a mitochondria and chloroplasts,
producing a complete new feature; however, two
processes would be important to complete phagocytosis.
First, the organism would have to disable its rigid cell
wall, leaving a malleable plasma membrane that could be
modulated to nd and surround a prey.
Second, the organism of course needed a mechanism
for projecting the membrane in a way that could easily
engulf its prey. This would require a cytoskeleton capable
to produce specic forces to open and close barriers. So
even if eukaryotic actin cytoskeleton was able to generate
a force on the membrane, it seems other two mechanisms
were necessary, rst a projection force produced by
polymerization and second, a motor molecule to move
the actin laments and put them near each other or to a
membrane (Cox, Foster, Hirt, Harris, and Embley, 2008).
Under this idea, if all this was possible, then the
polymerization-based membrane protrusion process
would be able to develop almost naturally as a result
of actin assembly, while the add-on of contractile
machineries involving other steps to evolve the set of
motor molecules and actin-binding proteins (Wickstead
and Gull, 2011). The problem with this idea, is that some
archaea and mollicutes do not have cell walls (Cavalier-
Smith, 2002).
In this sense, eukaryotic actin-based process could
possible developed microlaments and tubulin- based
microtubules, because some of the laments of the bacterial
cytoskeleton are essentially cytomotive which mean
that they can produce movement without any assistance
from other proteins, so laments themselves can act as
linear motors pushed by the kinetics of polymerization/
depolymerization process. That’s why some researchers
have explained that in eukaryotes, this activity increased
the evolution of numerous classes of motors, as well as
nucleators, severing agents, tip-binding factors, and (de)
polymerases functions, while other cytoskeletal laments
appear to have a more indispensable function, offering
opposition to external force or acting as a support to the
cell (Wickstead and Gull, 2011).
Another hypothesis about how movement was
possible in cells is called the neomuran hypothesis,
which tries to explain the origin of archaebacteria and
its diversication. Cavalier-Smith (2002) explains it this
way: “Archaebacteria originated by two successive
revolutions in cell biology: a neomuran phase shared
with their eukaryote sisters followed shortly by a uniquely
archaebacterial one. The rst, neomuran phase was
an adaptation to thermophily and involved a really
major transformation of 19 key characters, including
replacement of the cell wall peptidoglycan murein
by N-linked glycoprotein and a great upheaval in the
cell’s protein-secretion and DNA-handling machinery.
The second, relatively minor phase of specically
archaebacterial innovations, notably replacement of
acyl ester membrane by isoprenoid tetraether lipids and
of eubacterial agellin by glycoproteins, involved further
adaptations to hyperthermophile and hyperacidity,
respectively. Substantially later, several lineages
independently readapted secondarily to mesophyll.
Lateral transfer of genes from the immensely older and
far more diverse eubacteria often played a role in these
secondary returns to mesophyll and may also have
done in the origins of archaebacterial hyperthermophily,
sulphate reduction by Archaeoglobus and methano-
genesis. Under this perspective, the origin of the rst
eubacterial cell could be 3700 million of years ago, with
peptidoglycan walls and photo- synthesis, and the origin
about 850 My ago of the ancestral neomuran cell, when
N-linked glycoproteins replaced peptidoglycan and the
pre-eukaryote neomurans evolved phagotrophy, internal
skeletons and the endomembrane system” (Cavalier-
Smith, 2002, p: 17).
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
Cuadernos de Neuropsicología
Panamerican Journal of Neuropsychology
With this in mind, cell origins have been explained
with other two major models; rst a fusion model where an
endosymbiosis event distributing the mitochondrion came
very early, or a fusion later model where endosymbiosis
happened after development of several intracellular
structures. Although the second model places accent
a prerequisite for phagocytosis-like mechanisms to be
present to facilitate endosymbiont acquisition which
is considerate the origin of the eukaryotic cell and
represents one of the fundamental evolutionary changes
in the history of life on earth (Gray, 2012).
Over time, the host archaeon enlarged its area to
relate with symbiont (without phagocytosis) to obtain
these superuous products. At that point, the host–
symbiont coordination could exist in anaerobic and
aerobic environments (Stairs, Leger and Roger, 2015).
This proto-eukaryote had an archaeal cytoplasm and a
hydrogen- produced an organelle also capable of oxygen-
dependent respiration. Later, after the major lineages
of extant eukaryotes varied from the last eukaryotic
common ancestor (LECA), and aerobic and anaerobic
metabolisms were differentially absent in anaerobic
and aerobic lineages, generating the variety of energy
metabolism and the present-day mitochondrion-related-
organelles (Martin, Müller, 1988).
At this point all these hypothesis, land on another
ingredient to allow motility in cells, this is the structural
and architectural properties of the cytoskeleton. So, it is
important to dene that the cytoskeleton is mainly contained
into three polymer systems: actin laments (Wickstead
and Gull, 2011), microtubules, and intermediate laments.
Actin laments have a long shape and are formed by thin
bers. They have about 8 mm in diameter and are the
thinnest of the cytoskeletal laments, and they are also
called microlaments. On the other hand, other types are
microtubules, and they participate in a wide variety of cell
activities, because they are protein motors that use the
energy of ATP to provide the motion to cell. Lastly, exist
intermediate laments, that are small and dependent on
substrate stiffness and indentation depth, their principal
function is structural (Lodish, Berk, Zipursky, et al., 2000),
to reinforce cells and to organize cells into tissues and
epidermal cells, which are composed largely of proteins
(Jalilian, Heu, Cheng, Freittag, Desouza, Stehn, Bryce,
Whan, Hardeman, Faith, Schevzov, Gunning, 2015).
Actin protein and its role in the cytoskeletal support
Talking specically about the actin cytoskeleton
function is worth to say that this is regulated by a plethora
of actin binding proteins and specic signaling pathways.
It is also controlled by a convoluted collection of over 15
diverse sorts of actin lament arrangements, which have
been identied in metazoans and can literally being modify
in both spatial and temporal intracellular distribution in
response to physical and environmental stimuli (Lodish,
Berk, Zipursky, et al. 2000).
Something remarkable is the fact that two lament-
forming protein families, tubulin and actin 7, dominate
the cytoskeletons of all eukaryotes (Satir, 2016). From
a microscopical perspective, actin laments are semi
exible polymers with Lp ~17 µm. They have a diameter
of ~7 nm, and they are constructed from dimer duos of
globular actin monomers, with a polar functionality; this
means that they have a fast and slow growing individual
end (they are called the plus end and minus end
separately). The minus end has a critical actin monomer
concentration that is ~6 times higher than that the plus
end (~0.6 μM and ~0.1 μM at the minus and plus end
individually). When the end of an actin lament is exposed
to a concentration of monomeric actin that is above its
critical absorption, the lament end binds monomers
and grows by polymerization (Satir, 2016; Lodish, Berk,
Zipursky, et al. 2000).
This mechanism is important because contrariwise,
when the concentration of monomeric actin is below the
critical absorption, monomers separate from the lament
end, and the lament shrinks by depolymerization.
Basically, by having these two different critical actin
concentrations at the opposing ends of the lament, actin
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
Cuadernos de Neuropsicología
Panamerican Journal of Neuropsychology
laments can ourish asymmetrically, so when the actin
monomer concentration is between the two values, only
the plus end matures while the minus end shrinks, in a
back and forth dancing. This process, when the stretch
of the lament stays nearly constant and the polymerized
monomers inside the actin lament transfer motion
forward due to asymmetric plus end polymerization, is
known as tread milling, and this is a critical feature of how
polymerizing actin laments can generate force (Medina,
Worthen, Forsberg, Brenman, 2008).
At the same time, it is worth to say that microtubules,
are the strongest of the biopolymers, with Lp ranging
from 100 to 5000 µm depending on the lament length,
(Hightower and Meagher, 1986), and act as spirals
that may be rmly packed into packages where all the
helices are associated, and this arrangement is critical
to movement process (Satir, 2016). After all, actin is a
globular component of the cellular cytoskeleton and one
of the most abundant cellular proteins in cells (Jalilian,
Heu, Cheng, Freittag, Desouza, Stehn, Bryce, Whan,
Hardeman, Faith, Schevzov, Gunning, 2015), and the
best conserved eukaryotic protein (Satir, 2016) found
from unicellular organism to plants, animals, (Siccardi
and Adamatzky, 2016; Medina, Worthen, Forsberg,
Brenman, 2008; Hightower and Meagher, 1986) and
fungi (Roy-Zukav, Dyer, Meagher, 2015), so certainly the
mechanisms involved and highly mature.
Of course a high level of efciency mechanism is not
easy to design, so it is not a surprise that 60 actin-binding
proteins approximately, have been described in animals
and of course, contribute in a hug number of vital cellular
processes, such as cytoskeletal structure, conservation
of cell shape, cell motility, cell division, endocytosis and
intracellular transport, (Guljamow, Delissen, Baumann,
Thünemann, Dittmann, 2012), vesicle and organelle
movements, cytokinesis, muscle contraction, (Goodson
and Hawse, 2002) modulation of a variety of membrane
responses, translation of several mRNA species, and
modulation of enzyme activity and localization within the
cell (Monshausen and Haswell, 2013).
So, we can say that actin is a member of a larger
superfamily of proteins (Thomas and Staiger, 2014),
which acts as an expressway connecting diverse points
of the cell applying molecular motors driven by lament
assembly energies to transport proteins and organelles
across the cell’s limits (Yi, Huang, Yang, Lin, Song,
2016). As addition, the polymerization into laments is a
remarkable characteristic, which is the basis of functional
adaptability as result of an wide prevalence of actins in the
living world (Bertola, Ott, Griepsma, Vonk and Bagowki,
Another important characteristic about actin, is the
fact that is essentially regulated during cell migration, cell
adhesion, cell division, and several other essential cellular
functions, because actin is part of the conguration of
many cellular structures including lopodia, lamellipodia,
microvilli and stress bers (Zhu, Zhang, Hu, Wen and
Wang, 2013).
Such level of regulation is possible thanks to a
network design. Eukaryotes employ additionally more
than 100 actin binding proteins (ABPs), generally falling in
two classes with either actin monomer or lament binding
properties. The several interactions of ABPs with actin are
believed to be dependable for the evolutionary limitation
on its arrangement, making it one of the best conserved
proteins (Van den Ent, Amos, Löwe, 2001). Excluding
conventional actin, eukaryotic cells similarly contain actin-
like (ALPs) and actin-related proteins (ARPs), which have
well-characterized functions in cytoskeletal processes
(Venticinque, Jamieson, Meruelo, 2011).
Until now, six primary actin isoforms have been
recognized in superior vertebrates, (Goodson and
Hawse, 2002) and arthropods, (Brunet and Arendt,
2016; Monshausena and Haswell, 2013). being alpha-
skeletal (ACTA1), alpha-cardiac (ACTC1), alpha-smooth
muscle (ACTA2), gamma smooth muscle (ACTG2), beta-
cytoplasmic (ACTB) and gamma-cytoplasmic isoactin
(ACTG1). Moreover, actin can be organized in three pairs:
two isoforms expressed in striated muscle (skeletal and
cardiac tissue), two isoforms from smooth muscle (alpha-
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
Cuadernos de Neuropsicología
Panamerican Journal of Neuropsychology
smooth muscle predominately in vascular tissue and
γ-smooth muscle in the gastrointestinal and genital tracts)
and two cytoplasmic isoforms (Bertola, Ott, Griepsma,
Vonk and Bagowki, 2008; Murrell, Oakes, Lenz & Gardel,
As complex and it can look, actin exists predominantly
in one of two forms: monomeric actin (called G-actin)
and lamentous actin (called F-actin). The interaction
alongside these two actin systems is closely controlled
by a specic collection of proteins that bind actin directly
or indirectly. For example, Actin Depolymerizing Factor
(ADF), also known as Colin represents one actin-binding
protein that can strip actin by splitting and depolymerizing
actin laments (Bertola, Ott, Griepsma, Vonk, and
Bagowki, 2008; Van den Ent, Amos, Löwe, 2001).
While the actin-depolymerizing factor/colin (ADF/
CFL) gene family proteins have been associated in
cellular processes from membrane and lipid metabolism
to mitochondrial sustained apoptosis, a temporal-
specic dividing of expression arrangements proposes
that ADF/CFL protein variations have sub-functionalized
but might have gained different functions during their
evolutionary history. In this regard, some authors believe
that mammalian CFL and ADF/Destrin have biochemical
differences that are particularly recall functional
divergence (Roy-Zukav, Dyer, Meagher, 2015).
It is believed that “all living beings are in fact
descendants of a unique ancestor commonly referred to as
LUCA (the Last Universal Common Ancestor)” (Forterre,
Gribaldo, Brochier, 2005), even if it’s just a proposed life
system that apparently was the progenitor of the three
domains of life (Archaea, Bacteria and Eukarya), LUCA
probably can explain why motility become so important
to species.
In this regard, the eukaryotic cytoskeleton seems
to have evolved from ancestral precursors related to
prokaryotic FtsZ (which is a protein encoded by the ftsz
gene) and MreB (which is a protein discovered in bacteria
that has been documented as a homologue of actin) that
show 40− 50% sequence mainly across different bacterial
and archaeal species, meaning after million of years is
still around (Wickstead and Gull, 2011).
It seems that FtsZ is a plastid-derived and have
a similar role in the division of the chloroplast and/or
mitochondrion as in previous their free-living ancestors. So,
it is assumed that FtsZ mediates prokaryotic cell division,
and mitochondrial and plastid division in eukaryotes, by
developing an energetic ring among potential daughter
cells (or daughter organelles) (Koumandou, Wickstead,
Ginger, van der Glezen, Dacks and Field, 2013).
Before cytokinesis, which is the physical progression
of cell division, distributing the cytoplasm of a parental
cell into two daughter cells, admitting two types of nuclear
division called mitosis and meiosis. So, it is important to
notice that mitosis and each of the two meiotic divisions
result in two separate nuclei contained within a single cell
(Cooper, 2000).
It’s just a theory that cytoplasmic division of a cell
was able to create mitosis and meiosis, deriving the
common ancestor, so FtsZ was distributed to bacteria and
euryarchaeal, but since it is found in almost all modern
species and shows surprising plasticity in composition,
with the core lament-forming proteins conserved in all
lineages, the idea is highly suggestive (Forterre, 2005).
For the most part it is believed that FtsZ was also
used for division in the youngest eukaryotic and later, it
was involved as an actin-based machine for cytokinesis,
and eukaryotic FtsZ experienced a radical change and
it evolved into tubulin. Cytoskeletal proteins perhaps
evolved even earlier, in the common ancestor of bacteria,
archaea and eukarya, but FtsZ in particular is considerate
an ancient protein, because FtsZ and MreB (which is a
protein found in bacteria and identied as a homologue of
actin) (Koumandou, Wickstead, Ginger, van der Glezen,
Dacks and Field, 2013; Wickstead and Gull, 2011; Cox,
Foster, Hirt, Harris and Embley, 2008), and it has seen
that even ciliates contain actin, although ciliates are
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
Cuadernos de Neuropsicología
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microbial eukaryotes with two types of nuclei: a germline
micronucleus (MIC) and a somatic macronucleus (MAC)
(Faguy, Doolittle, 1998).
Of course, another important process that has
had a very long evolution, is the control of actomyosin
contraction, produced by an increase of intracellular
calcium, which is a well-preserved mechanism capable
to create mechanical stress in animal cells and underlies
muscle contraction, cell migration, cell division and tissue
morphogenesis (Poole, Lundin and Rytkönen, 2015).
Much of this complexity evolved before the last common
ancestor of eukaryotes, meaning that probably, the
distribution of cytoskeletal laments situates limitations on
the likely prokaryotic line made possible eukaryogenesis
process, which is estimated to have occurred over one
billion years ago (Wickstead and Gull, 2011).
It is hard to ignore that this crucial process in animal
muscle physiology is an ancestral feature of eukaryotic
cells (Tekle and Williams, 2016). However, it was necessary
ATP to promote the rotor stator-type ATPase, explained
before, so a protein that is as universal among cells as the
code, and no doubt is an invention of the world of genes
and proteins. After that, probably as Forterre (2005 p.
797) explains: “RNA played both the role of catalyst and
genetic material and this could happen through several
steps. After that, a new kind of cell began to have different
needs while interacted with environment and eventually;
actin was needed to allow new sets of skills”. As a result,
proteins as actin family and genes can be found within all
phylogenetic trees, and some analyses show that actin
genes could be divided into two major types of clades:
orthologous group versus complex group. Codon usages
and gene expression arrangements of actin gene copies
were stable among the groups because of basic functions
needed by the organisms but diverged within species
due to functional diversication. In this sense, most
vertebrates hold two genes for class IX myosins while
in invertebrates, a single gene for class IX myosins has
been classied. The two class IX myosins in mammals,
myosin IXa (Myo9a, myr7) and myosin IXb (Myo9b, myr
5), subsist in diverse variations among species (Newman,
Since actin, myosin and calmodulin are virtually
universally present in eukaryotic genomes, is important to
study them separately. Myosin is constituted by a heavy
chain containing the motor domain converting ATP-
hydrolysis into mechanical energy along actin lament
(with ATPase and actin-binding activities) (Newman,
2016), and usually a light-chain binding neck domain.
In most myosin families, the light chains are calmodulin
proteins; in others, specic calmodulin-related proteins
have evolved, such as the essential and regulatory light
chains of myosin II, while calmodulin is involved in the
regulation of a number of intracellular processes, including
cell proliferation (Luciano, Agrebi, Le Gall, Wartel, Fiegna,
Ducret, Brochier-Armanet, Mignot 2011).
In other species, such as vertebrates, cytoplasmic
actins look-like actins are present in many amoebas,
yeast and slime molds, this is because invertebrate
muscle actins are associated to vertebrate cytoplasmic
actins more than to vertebrate muscle actin isoforms. It
seems than actin isoforms particularly for striated muscle
tissue rst evolved in primitive chordates (Newman, 2016).
Talking about early amphibians or stem reptiles, this gene
maybe duplicated, which resulted in an alpha-skeletal
and a modern alpha-cardiac isoactin. The smooth muscle
isoactins are assumed to evolved during later development
of warm-blooded vertebrates and likely originated from an
early skeletal muscle actin. So far, over 30 different actins
have been dened from diverse muscle sources, some of
them are known by having a very specialized role (Faguy
and Doolittle, 1998).
In this sense, when eukaryotic cells began to change
external stimuli into membrane depolarization, and turn
on triggers effector reactions, such as secretion and
contraction, permitted to convolute a number of important
and diverse cellular processes such as organelle
movement, exo and endocytosis, nuclear transporting,
and chromatin repair, so a variety of classes of actin
binding proteins are found in plants and animals that
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facilitate the vigorous nature that makes it one of the
most dynamic characteristics in a eukaryotic cell (Murrell,
Oakes, Lenz & Gardel, 2015).
So far, we can say that actin proteins family has a
very important function to movement process, intra, extra
and among cells. Being a very well conserved heritage
from prokaryotes cells, there is no doubt that is a relevant
part of the evolution of species. But, how did this happen?
Evolution of cells and movement processes
It accepted that life rst began at least 3.8 billion
years ago, around 750 million years after Earth was
formed. Some theories claim that the rst cell started
by the insertion of self-replicating RNA in a membrane
composed of phospholipids. It is known that these are
the basic components of all the biological membranes,
including plasma membranes in both prokaryotic and
eukaryotic cells (Cooper, 2000), and ho we have been
exposing this lead to a dynamic strength for the evolution
between prokaryotic, bacterial, archaeal, and eukaryotic
cellular organization (Brunet and Arendt, 2016).
However, because cells required energy to move, the
mitochondrion was likely the best mechanism possible,
and it is well known for its function in ATP synthesis by
oxidative phosphorylation. In this process, pyruvate
from glycolysis is imported into mitochondria where it is
oxidatively as decarboxylated to acetyl-CoA by Pyruvate
Dehydrogenase (PDH) and becomes part of the Krebs
cycle to produce nicotinamide adenine dinucleotide
NADH, and Flavin adenine dinucleotide FADH2, its
function is to provide electrons to the electron transport
chain. They both transport electrons by exchanging a
hydrogen molecule to the oxygen molecule to produce
water during the electron transport chain; these reduced
cofactors link chemically with oxygen, by the electron
transport chain (ETC), to produce a proton gradient across
the inner mitochondrial membrane and nally reduce O2
to H2O (Stairs, Leger and Roger, 2015).
At the same time, the proton force drives ATP
synthesis by an F1Fo-ATP synthesis. However, in addition
to holding genomes that are replicated, transcribed and
translated, mitochondrial process has an important
function developing iron–sulfur (Fe–S) cluster generation
(via the iron–sulfur cluster (ISC) system) as a biosynthesis
process, since amino and fatty acid, phospholipid, vitamin
and steroid metabolism are necessary to cells (Newman,
Given these points, in 1998, Martin & Müller, proposed
the hydrogen hypothesis in which they discussed
if eukaryotes could have risen across the symbiotic
relationship of an anaerobic, strictly by hydrogen-
dependent, strictly autotrophic archaebacterium (the
host) with a eubacterium (the symbiont) that was able to
breathe, but capable to produce molecular hydrogen as a
waste product of anaerobic heterotrophic.
In this regard, these authors explain the host’s
dependency upon molecular hydrogen, created by
the symbiont, as the selective source that put-on the
common ancestor of eukaryotic cells in motion. With
this development, it is believed that the ancestor of
mitochondria was an H2-product, mainly by an anaerobic
a-proteobacterium that had a syntrophic relationship with
a hydrogen-dependent methanogenic archaeon, however,
in an anaerobic ecosystem, the a-proteobacterium created
ATP by the anaerobic extended glycolysis pathway,
generating hydrogen, and of course it was necessary
carbon dioxide and acetate as discarded products that
were spent by the methanogen (Martin, Müller, 1988).
The selective benet of these changes was the
ability to remain producing acetyl-coenzyme A and
eventually ATP from pyruvate (and/or malate) under
hypoxic conditions usually encountered by free-living
and anaerobic eukaryotic systems. However, with a
need to adapting to new atmospheres, eukaryotes could
acquire and express genes from prokaryotic or eukaryotic
donors that permitted them to succeed (Poole, Lundin
and Rytkönen, 2015), since it has both molecular and
morphological attributes very conserved, they have
participated as an essential role in the understanding of
the origin and evolution of different eukaryotes (Tekle and
Williams, 2016).
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Besides the mitochondria, of course centrosomes
are another old improvement. They are membrane-free
organelles that serve as main microtubule systematizing
cores in different eukaryotic lineages (Azimzadeh, 2014).
In preparation for cell division, the centrosome duplicates
during mitosis, so the sister centrosomes act as an
important way to determine the indispensable bipolarity
of the spindle. Because the role of mitosis is to divide a
mother cell into two genetically identical daughter cells,
the cell must guarantee that the centrosome inherited
from the previous mitosis doubles once and only once
(Sluder, 2014).
Such strategies are just an example of the multiple
survival processes that evolution created with a range
of delightful movement options. This is particularly
interesting if it’s seen in perspective, especially when we
think that animals (Metazoa) are just one of some dozen
freely developing groups of multicellular organisms. It is
believed that they emerged more than 600 million years
ago, including cells belonging to a bigger phylogenetic
group, holozoa, which also involves some existing
unicellular and transiently colonial systems (Tekle and
Williams, 2016).
While plants, bacteria and virus are modest
examples of motility, the transition from few cells organisms
to vertebrates in water is a fundamental step in the
evolution of terrestrial life, and the exponential expansion
of bones and muscles became a very necessary item.
Once animals left the aquatic environment, required a
skeleton capable to resist the signicant effects of gravity
for example, as well as permit operative conduction of
force to the substrate to allow propulsion, so it is not a
surprise that in most terrestrial vertebrates, the bones of
the appendicular skeleton provide this framework (Blob,
Espinoza, Butcher, Lee, D’Amico, Baig, Sheffeild, 2014).
As an example of this, a variety of actinopterygian sh
species evolved to acquire the ability to navigate over land
using combinations of ns that are prolonged by exible
bones. Some critical improvements to this evolution was
the development of a weight bearing pelvis, hind limbs
and their related musculature and movements that allow
running or walking back and forth, and probably this
feature allow them to dominate in terrestrial locomotion.
The fossil record exposes how the skeletal structure of
the load-bearing limbs of tetrapods (animals descended
from sh) has developed, but since soft tissues are not
usually conserved as fossil evidence, so there is not
clear evidence of how the dramatic alterations of the limb
musculature started to change (Blob, Espinoza, Butcher,
Lee, D’Amico, Baig, Sheffeild, 2014).
No to mention that locomotor strategies in terrestrial
tetrapods have evolved from the use of sinusoidal
retrenchments of axial musculature, ostensible in
ancestral sh species, to the dependence on powerful
and complex limb muscles to provide propulsive force,
this means the implementation of the fully derived mode
of hind limb muscle formation from this bimodal character
state is an evolutionary innovation that was critical to the
accomplishment of the tetrapod transition (Cole, Hall,
Don, Berger, Boisvert, Neyt, Ericsson, Joss, Gurevich,
Currie, 2011).
Nevertheless, even if muscles, nerves and
somatosensory processes are a big leap in evolution
terms, a central nervous system was necessary to
generate adaptive strategies. The origin of the nervous
system was an evolutionary event that essentially changed
how control is achieved within a multicellular body.
Nervous System: controlling the movement process
It is vastly accepted the assumption that the human
brain weights in average 1.2–1.8 kg, and has around 100
billion neurons (Jékely, Kejzer and Godfrey-Smith, 2015).
Although, it is believed that the origin of brain and central
nervous system (CNS) can be marked by the Paleozoic
era, 540 million of years ago (Strausfeld and Hirth,
2016), At the same time, it’s believed that the origin and
diversication of the animals occurred throughout the so-
called Cambrian explosion, during a period when many
important organ systems appeared (Kass, 2013). In this
sense, the nervous system of humans must be considered
the best draft in nature, not only among animals, but from
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sponges, arthropods, chordates and placozoans (Budd,
As result of such a progress in nervous systems, early
mammals developed from mammal-like synapsids over
200 million of years (Hejnol and Lowe, 2015). Synapsids
are the dominant large terrestrial animals in the world,
so they conquered the oceans like whales, pinnipeds
and the air, for example bats, however they differed from
mammals, as Kass (2008) explains, because they had
“low-resolution olfaction, poor vision, insensitive hearing,
coarse tactile sensitivity, and unrened motor coordination,
together with limited sensorimotor integration”
By the same token, evolutionary research has
explained how such level of differentiation occurred, for
example early mammals had tiny brains in comparison
to their body size, at the same time, they exhibited
considerably bigger forebrains, prominently expanded
olfactory (piriform) cortex, a dorsal cap of neocortex,
an expanded cerebellum, a thicker spinal cord, however
these brains controlled simpler sensorimotor systems.
There was also a good draft to auditory adaptations that
would warrant high frequency hearing, and perhaps they
could use high frequency communication calls, but later,
mammals emerged from mammal-like reptiles about 200
million years ago and radiated into the over 3,500 living
species (Collin, Davies, Hart and Hunt, 2009).
For this reason, some researchers believe that the
relationship between distantly linked animals during
the development of their central nervous system could
lead the enlargement of a central nervous system with
a distinct centralized medullary cord and a subdivided
brain, since this is homologous across bilaterians, and
then a morphologically and molecularly tri-partitioned
brain connected to a central nervous system was
developed in the nal common ancestor of protostomes
and deuterostomes, such idea also suggests a reduction
in animals that have a much simpler organization of their
nervous system (Bielecki, Høeg, Garm, 2013).
Important to realize, is that from a phylogenetic
perspective, the typical debate of the origin of the nervous
system is whether or not it had one or more distinct origins.
Yes, it is believed that nervous systems evolved once only,
at the base of the so-called Epitheliozoa basically all of the
animals separately from the sponges. However, the best
indication for early nervous system remains the Ediacaran
to Cambrian as a fossil record, but its complexity across
species cannot be understood as increasing nervous
system development, because an ecological aspect also
seems to play a role in determining trace fossil morphology
(Kass, 2013).
Under this context, it seems that motility has a
distinct impact in human development (Dzib-Goodin, and
Yeliza rov, 201 6), and spec ies ev olut ion, not on ly fro m a
genetic, and cultural perspective, but also as developing
cause to advance as specie, since movement is related
with processes such as learning, memory and sleep,
through many neurological networks shared for all these
systems (Lotem and Halpern, 2012; Dzib-Goodin, Sanders,
Yeliza rov, 2017). Th es e as so ci ation s ar e im po rt an t to
cognitive skills and learning in order to help species to
adapt to the environment.
Additionally, when we talk about the central nervous
system, there are some considerations about the origin of
sensory organs and how they could be important to brain
evolution and movement process. In this regard it can be said
that the origin of eyes for example, has dominated debates
and theories about what selection forces have driven eye
evolution; so it can be said that it was more than 540 millions
of years ago when the rst appearance of photosensitive
receptors as single lens eyes and multifaceted eyes and
their underlying circuits, not to mention that color vision
evolved in the earliest vertebrates, providing the source
for color perception in all extant vertebrate classes found
today (Bielecki, Høeg, Garm, 2013).
With this in mind, it is important to understand the
evolutionary limitations placed upon the shape, light
reactions, spectral sensitivity and molecular assembly
of photoreceptors in early vertebrates and their role in
visual behavior, because paleontological evidence from
the Silurian and Devonian periods shows that the lateral
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eyes of the ancestral vertebrates were skilled to create
image formation and rotate within their own orbits by seven
extraocular eye muscles (Perrin, Sonnemann, Ervasti,
This means if all sensory systems react due to
receptor adaptation, visual systems are not the exception.
Also, since photo adaptation happens at a cellular level of
photoreceptors it was an inevitable feature in metazoan
vision. Thus, since photoreceptors adapt to constant
visual stimuli and counterstrategies are necessary to
prevent image fading or blindness. The best-known mean
to prevent adaptation is the xational eye movements in
mammals (known as tremor, drift and micro saccades),
which unceasingly refocus and renew the retinal image.
These movements are produced by an oculomotor
system and since they have a blurring consequence on
the retinal image, additional neural adaptations in post-
processing pathways have evolved to avoid the interludes
of movement These mechanisms are very powerful, but
also very expensive in both energy and neural capacity,
so they are not available for animals with less elaborate
visual processing (Collin, Davies, Hart and Hunt, 2009).
Of course, it is not possible to forget the auditory
system, which seems to be exceptionally sensitive to
perturbations of cytoplasmic actins, possibly because
actin is a key physical component of auditory hair cells,
which transform sound waves to neural signals. Hair
cells are contained in the organ of Corti, both of which
feature a complicated architecture that is a requirement
for appropriate function. The organ of Corti consists of
three rows of outer hair cells and one row of inner hair
cells, organized with several types of support cells.
This ribbon-like structure goes longitudinally alongside
the length of the cochlea. External hair cells improve
sensitivity to sound, while inner hair cells are the auditory
receptors. Both cell types are crowned with specialized
structures called stereocilia, which are detailed microvilli
made from a mixture of b-actin and c-actin laments that
are organized in a strongly bundled para-crystalline array
(Chakraborty and Jarvis, 2015).
As a result of the specialized sensory motor systems,
the nervous system enhanced the level of diversication.
In this sense, it is worth to mention that the human–
chimpanzee divergence is commonly estimated at 5–6
million of years however, some researchers consider this
divergence could be greater than 7–9 million of years
(Kass, 2013), meaning brain motor system could be used
long before the human brain.
The primates brain evolution
There is no doubt that the density in skills and motor
coordination of human brain is a highlight moment among
primate’s evolution. Only the neocortex constitutes about
80% of the human brain, and this is segmented into diverse
specialized regions, this is only one reason that this brain
like that mediates accomplishments and abilities has no
comparison among any other species (Kass, 2013) and it
is the reason for a unique adaptation in motor skills (Dzib-
Goodin and Yelizarov, 2016).
Generally, the origin of nervous systems has
been judged through two different theoretical models.
Hashimoto, Ueno, Ogawa, Asamizuya, Susuki, Cheng,
Tanaka, Taoka, Iwamura, Suwa and Iriki, (2013) call
them the “input–output (IO) and internal coordination
(IC) models”. The two models highlight two distinguishing
features of the nervous system as a control device. These
authors explain that IO models, have the main function
to receive sensory information and process it to produce
meaningful motor output.
As a result of such specialization, there is a difference
regarding IC and IO roles, because they have two
different functions: behavior, and also physiological roles,
so it is possible to distinct three types of effectors that
the nervous system can affect these: are cilia, muscles
and glands. On the other hand, some physiological
processes involved internal organization. In this sense,
complex, muscle-driven physiological processes, such as
peristaltic spasms that move the content of the stomach
or heartbeat, require IC systems to switch them; while an
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IO model tends to adopt an operational effector system,
an IC model highlights the evolutionary change capable to
produce new multicellular effectors. In particular, the use
of large contractile tissues (muscle) by large organisms, is
an important evolutionary development, since movement
in a muscle is a challenging task that should not be taken
for granted (Jékely, Kejzer and Godfrey-Smith, 2015).
For example, ciliary pounding can be used for
propulsion in an extensive range of small systems, but
also has other uses. For instance, inside a sponge, cilia
are used to allow water ow to permit access to food and
oxygen, so the cilia must have coordinated movements,
and this is a scenery in which an IC function inuence
can be relevant. Once a coordinated ciliary signal exists
in an organism, other control strategies may adjust the
activity of the cilia. Subsequently, cilia can become part
of an IO system. In this sense a phototactic routing is an
important IO function that is explicit to locomotion and can
be found even in many metazoan larvae (Jékely, Kejzer
and Godfrey-Smith, 2015).
With a much more complex system, the brain of the
genus homo could be developed in the early Pleistocene,
just after 2 million of years. But long after, around 200
thousand years ago, the rst draft of the Homo Sapiens
can be recognized in fossil records. However, it’s important
to mention that the lack of fossils related to this period
makes interpretation difcult, however some evolutionary
patterns can be considerated, for example the pelvises
of early Homo, are similar in general about the shape to
earlier hominids, and have traits that differentiate them
from australopithecines, and it can be said that many
of these traits are perhaps linked to modications in
locomotor performance (García-Grajales, Jérusalem,
Goriely, 2017).
This is because in order to control so many new
characteristics, the nervous system had to suffer an
adaptive environmental inuence. Since neuronal growth
is a key process necessary to establish the neuronal
network during neurogenesis, this could be consequence
of all the new requirements that environment was
demanding from brain. Besides its fundamental role,
neuronal growth also balances critical needs in human
brain plasticity and neuronal renovation during all cycle of
life (Kass, 2012a).
In order to warranty the process, numerous neurites
from the soma, produce a highly dynamical hand-shape
extensions called growth cones, this is a self-care process
to human brain, and it can be found since early stages
of neuronal development. This will continue growing,
until one neurite specializes hooked on the axon, while
all the other neurites become dendrites. This process
was probable thanks to Paralemmin-1, which is a protein
that stimulates cell development in plasma membrane. A
family of these proteins can be found on vertebrates, and it
has been possible the identication of paralemmin genes
in the different vertebrate genomes, so it is believed they
have a common gene organization (Khaitovich, Weiss,
Lachmann, Hellmann, Enard, Muetzel, Wirkner, Ansorge,
Pääbo, 2004).
The impact on this can be explained as a result of the
changes in motility in neurons, particularly long neurites
packed with G-actin need control the development of
F-actin in reaction to dynamic events such as synapse
structure or axon regulation through sensation of chemo-
attractive/chemo-repulsive signals. Another key point is
that the formation of ectopic F-actin need to be blocked
to prevent physical obstacles that might obstruct vital
transport roles inside these thin neurites and create
damaging cellular results (Kass, 2004).
For example, it is known that neurites include a
microtubule-rich cytoskeleton that offers a physical
support to delivery both inside and outside directions for
cargoes necessary to keep correct neuronal functions.
So, in this regard it is necessary an energy-dependent
molecular motor, including dyneins and kinesins, which are
ATPases that physically assist sending targeted cargoes
by directional motion lengthwise these microtubules.
The kinesin superfamily protein KIF5 in particular is
able to transport various cargoes involving membranous
organelles, cytoskeletal proteins, and mRNAs (Khaitovich,
Weiss, Lachmann, Hellmann, Enard, Muetzel, Wirkner,
Ansorge, Pääbo, 2004).
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This is particularly important to neurons, since
once an axon found an area to stablish, it will have to
spread through spaces travelling across a rich chemo-
mechanical signals, but it won’t escape of complications
until is able to locate its nal reach-point, but physical
forces will be acting, from molecular structures of the
neuron organelles to the nal formation of the whole
organ. So, it is signicant to notice that the main physical
framework of the neuron, the cytoskeleton is an evolving
active polymeric association that is actively participating
in axonal outgrowth during a long period of time, in human
brain (Kass, 2004).
Previously it was mentioned the F-actin and G-actin.
Since the cytoskeleton is a product of evolution, this is
constituted by three main kinds of lamentous polymers:
F-actin, microtubules and neuro- laments. Neurolaments
are inactive and apolar polymers. Although are the most
copious cytoskeletal laments in the axon, it seems they
do not contribute during axonal growth. In contrast, the
two other polymers, F-actin and microtubules, are really
dynamic and polarized. “The old polymerizes at one end
(barbed-end) by addition of G-actin and depolymerizes
at the other end (pointed-end) by removal of monomers,
while the latter polymerizes at one end (plus-end) by
addition of tubulin dimers and depolymerizes at the
other end (minus-end) by removal of monomers. While
microtubules are the rmest cytoskeleton components
and F-actin are less rigid on their own, the latter are able
to build organized stiff structures thanks to the presence
of high concentrations of crosslinkers. Their complicated
interactions as well as their relations with the surrounding
structures and associated motor proteins (e.g., Dynein
or Kinesin for microtubules or Myosin II for F-actin) are
crucial for proper axonal development, they also are
heterogeneously dispersed along the axon domain”
(Wickstead and Gull, 2011, p 515).
This gives to the cytoskeleton the capacity to respond
is such vigorous way both mechanical or chemically
and allow the development of so many arrangements,
structures and skills that permit cells to performance
the way they need into a specic background, during
growth and renovation as a key process in development
of species and evolution in a higher standpoint (Dzib-
Goodin and Yelizarov, 2016).
Through molecular motors, the cytoskeleton is able
to get energy from ATP hydrolysis, transforming it into
mechanical energy that can provide energy to the system
into arrangements produced with not thermal motion
alone. Beside with the characteristic shape of cytoskeletal
laments, which can assemble or disassemble quickly
with chemical species gradients or regulatory signaling
cascades, that allow to this nature item to respond in such
particular way to the needs of cells (Popov, Komianos,
Papoian, 2016).
However, it can’t be denied that different anatomical
brain structures developed at diverse times during
vertebrate evolution, based on different needs of the
environments and of course thanks to the new designs
possible depending the kind of motion needed, and this
produced the vertebrate brain known by its three divisions,
with the spinal cord and brainstem (hindbrain, midbrain
and thalamus) having more preserved constitution,
maybe because it adapted to more dependable skills, and
the telencephalon with a more varied organization, which
exhibit three major structures, the pallidum and striatum
having more well-preserved organization, and the pallium
or cortex, with a more different organization. While the
pallium is primarily hidden in mammals, it is typically
nuclear in birds, reptiles and other vertebrates, mainly
because needs over the environment are different in
every specie. However, some changes happened with the
appearance of the telencephalon through the invertebrate
to vertebrate evolution, because diverse motions were
required, denoting that the central nervous system has
been an central target of selection (Khaitovich, Weiss,
Lachmann, Hellmann, Enard, Muetzel, Wirkner, Ansorge,
Pääbo, 2004).
At a molecular level, even if it has been accepted that
more of those changes are due to Darwinian selection,
that perspective was challenged by Kimura’s neutral
theory of molecular evolution (cited by Khaitovich, et. al,
2004). This theory is based on the vast differences seen in
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nucleotide and amino acid sequences within and between
species, that have no or only minor selective results. So
now it seems, their incidence within a species and the
xation of differences between species are mainly the
consequence of stochastic processes, meaning that are a
collection of random adaptations (Kass, 2004).
This could happen in the middle of adaptions and
deviation of species primates appeared around 80 million
years ago, as a branch of the Euarchontoglire superclade.
Kass (2004) explain that: It is believed they were short,
arboreal, and nocturnal creatures; they fed on small insect
and vertebrate prey, buds, and fruit Primates constitute an
order of mammals that is extremely diverse in brain size”.
This branch particularly covered abundant lines of archaic
primates that found extinction, and the branch euprimates
that began to the current galagos, lorises, tarsiers, and the
greatly varied anthropoid monkeys, apes, and hominids
(humans and extinct species more closely related to
us than chimpanzees), so they are in a considerate our
common ancestor.
When Kass, (2012a) explains about the brain
characteristics of this lineage, he says those brains were
slightly stretched, but they don’t have a big size and they
have a similar proportion to body size than the brains of
extant prosimian primates (lemurs, lorises, and galagos).
“Their eyes were large, and frontally directed, and their
temporal cortex was enlarged”. Consequently, it means
that vision was important to survive in the environment,
modications for life in these branches of trees suggested
that their neural systems for eye-hand coordination were
well established to jump from tree to tree (Kass, 2012a).
It is accepted that the closest living ancestors
of primates are the Scandentia (tree shrews) and
Dermoptera (ying lemurs) of the Archontan branch of
Euarchontoglires”, while the more distant are the “Glires
branch includes rodents and lagomorphs”. Although
humans and chimpanzees are unrelated from a common
ancestor by a few million years, human brains are three
times bigger, and had maximum of that increase over
the past 2 million years of hominin evolution. Only within
thousands of years, human survived to their relatives,
mainly because a great ability to move, create with their
hands and think. The youngest disputed hominin is the
Sahelanthropus tchadensis, who lived approximately
7 million of years ago; so, there is no doubt that the
emergence of the homo erectus sensulato in East Africa
characterizes a fundamental turning point in hominin
evolution (Maslin, Schultz and Trauth, 2015), and we still
see their adaptation to the environment.
In this regard, the relative expansion of the cerebellum
in primates besides to stereopsis and amplication of
the visual coordination apparently reinforces primates
ne viso-motor control and manual dexterity. This was
particularly important, to search fruits and probably hunt,
so this smooth-pursuit eye-movements in primates create
a unique cortico-cerebellar pathway that evolved at the
same time of foveal vision. “All major cortical regions, for
example beyond motor cortex and including frontal and
prefrontal areas, have reciprocal connections with the
cerebellum” (Kass, 2013), giving a more precise movement
process to stay alive and respond in the environment.
That’s why Kass, (2012b) writes about: “these cortico-
cerebellar loops form multiple, independent anatomical
modules which are architecturally quite uniform”. And with
such design, it was opened the opportunity to other ne
movements, since generally speaking, some of the tools
can be used for a different task. The best example of this
is language process (Galván-Celis, Pechonkina, Slovec,
Dzib-Goodin, 2015). The reason for this is that the unit of
brains, the neuron, are not developed for a small or big
brain specically, but they will adapt their numbers and
structure depending environment needs. So, reasonably
rather than small brains developing small neurons, they
will have less, and bigger brains will have more neurons, in
this sense, additional growths in brain size could produce
less and less gain to analyze and use information. A
conceivable answer to why bigger brains become more
modular by aggregating areas and subdivisions of areas
in order to reduce the number of long connections
(Hoffman, 2014), is that there is a relationship between
energy consumption and the energy species get through
their diet, based on general activity.
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
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Of course, there were not only small other changes
that made the human brain a better natural draft among
species, lateralization for example, allowed to decrease
the prerequisite for huge amounts of long, dense axons
streaming through the two cerebral hemispheres motor
cortex (Mendoza, Merchant, 2014), another action that
must be added is the fact that in humans particularly,
movement process is more focused on a super specialized
digit movements, and of course, it means a specialization
of ventral premotor cortex of the left cerebral hemisphere
for speech (Khaitovich, Weiss, Lachmann, Hellmann,
Enard, Muetzel, Wirkner, Ansorge, Pääbo, 2004), since
this is a very elaborate skill.
This specialization, however didn’t began with homo
sapiens, early anthropoids showed many differences
from other primates, and part of the reason was a cultural
deviation, during 65–90 million years the diurnal niche
eating fruit, buds, and maybe insect in the deadly areas of
tropical forests, forced to an advance of posterior parietal
sensorimotor cortex that involved areas like visual, auditory,
and somatosensory in order to create a ne motor answer
to the environment, and it is hypothesized than the frontal
motor regions, as portions of a sensorimotor system,
were increased and subdivided in early primate (Kass,
2008), this allows for human brains to create a different
path and more specic and ne movements (Khaitovich,
Weiss, Lachmann, Hellmann, Enard, Muetzel, Wirkner,
Ansorge, Pääbo, 2004).
A good example of this process can be experimented
with a very curious phenomenon, when something is
touched with a single nger, can stabilize a person who
is potentially losing his/her balance. This means the
spatial perception of the ngertip is better distinct than the
vestibular system and it is efciently sensitive to detect
small body change. Maybe because tactile feedback from
the nger is basic for decreasing changing responses to
the environment, but no effects of ngertip-contact forces
on postural changes. An explanation is that bimodal
neurons in the vestibular cortex reach the vestibular and
somatosensory inputs might explain these effects on
vestibular responses. But at the same time vestibular
cortex inuence multimodal reference structures to
maintain the unity of the spatial experience as a recall
of the needs of the rst primates in a complete different
area, when they hand from trees (Barton, 2012).
That’s why is understandable that when Homo
Sapiens appeared, they were other motor behaviors
controlled by the nervous system, and culture began
to have an impact on the approaches to adapt to the
environment, so it opens the door to cognition processes
(Stout and Chaminade, 2012). So, it is logical to think
that using tools perhaps allowed more exquisite abilities
needed to survive, but undoubtedly is not the only one
reason, because some other mammals use tools to get
food, and by some reason didn’t forced to those species
to the level of human development.
Human brains: cognition and its relationship with
motor control
Speech and the use of tools are both goal-directed
motor actions (Stout and Chaminade, 2012). Now, the
classical description of the tool is limited to external
objects held by a hand in order to interact with the external
environments, but modern humans also use tools to
increase the reachable area or externalize our existing
sensory organs, or to support the detection of information
that is outside natural sensory range. This means that the
natural intransitive movement becomes transitive, and
this can create a “sense of the self (as the subject) and
leading to the movement of ourselves or our body parts
perceived as objects” (Iriki and Taoka, 2012).
But there are other tools, for example producing words
and vocal learning, are a critical component of spoken
language acquisition (Galván-Celis, Pechonkina, Slovec,
Dzib-Goodin, 2015), since they are dened as “the ability
to modify acoustic and/or syntactic features of sounds
produced, including vocal imitation and improvisation”
(Stout and Chaminade, 2012) and, similar than other motor
activities, this implies implementation and comprehension
of neural circuits integrating sensory perception and
motor control, so they are linked as a need to survive and
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communicate strategies, used into the environment (Iriki
and Taoka, 2012).
Certainly, can be controversial the idea of language
as a tool, since is easy to see a big difference between
speech and the way a tool is used, especially because is
clear that language is mainly a modality visual-auditory
and the use of a tool requires visuospatial, somatosensory
and manual skills, so the argument can be easily denied.
But there is a good counter-argument, anatomically
there are resemblances in the way speech and tool-use
networks are systematized, including strong evidence of
functional–anatomical intersection in inferior frontal gyrus
and in inferior parietal and posterior temporal cortex.
These discovers a similarity between cognitive processes
and cortical networks that use speech and tools, and this
explain why behaviors are best seen as special cases in
the more general domain of complex, goal-oriented action
(Stout and Chaminade, 2012).
Under this idea, it is plausible that the evolutionary
intensication of tool-use could incorporate the
combination of visual, and symbolic-abstract information
leading to the appearance of a novel functional brain area
for abstract understandings of tool functions, fullling
the condition for the boost of complex human tool-
usage (Hashimoto, Taoka, Obayashi, Hara, Tanaka, Iriki,
2013). This can be a good reason to explain why areas
of the neocortex are especially big in the human cortex,
for example the prefrontal granular cortex or language
related Broca and Wernicke areas, which are considered
as analyzers for integration of information from both
sensory and motor areas (Galván-Celis, Pechonkina and
Dzib-Goodin, 2014).
In this regard, Corballis (cited by Jablonka, Ginsburg
and Dor, 2012) explains that motor control necessary to
learning and teaching tool uses and fabrication, is the
scaffold for the increasingly complex communication,
emphasizing the role of motor control, arguing that the
evolution of language could be originated by the control
of manual and oro-facial gestures (and only later of
vocalizations). That why Corballis proposed that the
“voluntary motor control that was necessary for tool
making made gestural communication easy, and this was
generalized to oral movements, which then led to speech”.
Another process that is not easy to ignore is motor
imitation, which is a necessary ability to manufacture
complex tools, observed among the Acheulean, and some
think it was a prerequisite for the evolution of syntactic
language, this because as Iriki and Taoka (2012), explain
the recursive organization that adopts the combination of
motor units is essential to design complex tools and at the
same time, it is the basis of syntax, since message signs
are inserted and merged into semantic representations
giving order to every idea.
This is easier to understand because usually
language can be divided into a conceptual–intentional
system that deals with thoughts and meanings (Rakic,
2009), and a sensorimotor system that deals with the
acoustic analysis of speech sounds and their production
(Galván-Celis, Pechonkina and Dzib-Goodin, 2014), this
implies that once a original cognitive demand, such as
integration of motor tools into the body representation,
has become implanted in the environment, adjustments
of brain organization would be stimulated spontaneously
within the normal developmental processes in subsequent
generations. The incidence of such a plastic response
during the lifespan as a result of behavioral modications,
could be possible by the existing of an adaptive capacity,
and its subsequent consolidation (under selection acting
on changing gene frequencies), as a default state that is
unchanging over generations (Iriki and Taoka, 2012).
Eventually, other processes could be activated to
use the motor areas that have changed as an effect of
culture, and an example of this can be writing and reading
processes, since they are new learnings in the history of
humankind (Galván-Celis, Pechonkina and Dzib-Goodin,
Vocal control
Interestingly, vocal learning is a rare attribute into
other species, so far it is recognized in only ve remotely
related groups of mammals (humans, bats, elephants,
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cetaceans (dolphins and whales) and pinnipeds (seals
and sea lions) and three distant related groups of birds
(parrots, songbirds and hummingbirds). However, even if
evolved independently those lineages of vocal learning
birds and humans, both share clear forebrain pathways
that control the understanding and production of learned
vocalizations. In these pathways, all three avian lineages
contain seven cerebral (telencephalic) vocal nuclei and
several thalamic nuclei (Scharff and Petri, 2011).
These nuclei, are very well described in songbirds
and parrots, and they are distributed between two sub
pathways explained by Chakraborty and Jarvis. (2015)
these are ”1a): (i) the vocal production, or posterior,
pathway that inuences the production of learned song,
which includes an arcopallium nucleus (songbird RA
(robust nucleus of the arcopallium), parrot AAC (central
nucleus of the anterior arcopallium), hummingbird VA
(vocal nucleus of the arcopallium), analogous to the
laryngeal motor cortex (LMC) in humans that makes a
specialized direct projection to brainstem vocal motor
neurons (MN), which in turn controls the vocal organs,
the syrinx (birds) and larynx (humans); and (ii) the vocal
learning, or anterior, pathway that is primarily responsible
for vocal imitation and plasticity, which forms a pallial–
basal ganglia–thalamic loop”. This is equivalent to the
loops found in mammalian brains that include Broca’s
speech area in humans, specically ((Dzib-Goodin,
Yelizarov, 2016).
Something remarkable to mention, is the fact that the
song and speech regions in both pathways are inserted
in or adjacent to non-vocal motor brain areas, and these
non-vocal motor areas are present in other vertebrate
species studied and could be involved in the learning
process of non-vocal motor behaviors (Chakraborty and
Jarvis. 2015).
There is no doubt that in this evolutionary scenario
genes were important, and it has been said that the
expression of FoxP2, which is a Fork head box protein
P2 (FOXP2) is a protein that in humans specically, is
coded by the FOXP2 gene, and it is required for proper
development of speech and language. During the evolution
of vocal skills, once the striatum got attached to other
regions necessary for vocal learning to occur, FOXP2
mutated in humans and this might have affected neural
transmission, and in Area X of the striatum, consequently
became benecial for sensory motor integration or dened
timing of vocal gestures and to other motor learning tasks
in adjacent non-vocal circuitry cells (Scharff and Petri,
2011). This would be a two-hit consequence of FOXP2 ’s
role in language evolution, since if circuit changes, gene
function changes in consequence to adapt to the new
needs of the system (Galván-Celis, Pechonkina, Slovec,
Dzib-Goodin, 2015; Galván-Celis, Pechonkina and Dzib-
Goodin, 2014).
Tools manipulation
So, if language is linked with tools manipulations
and design, in a way to change gene expressions, it is
important to notice that primate manual manipulation,
including those on skilled human that are capable to
use a stone tool, have exposed three manipulative
abilities studied as unique to the human hand. The rst
is precision control, dened as the ability to rotate and
manipulate objects within one hand using the thumb and
ngertips. While other primates characteristically need to
use the palm as well or their other hand, a foot or the
mouth to manipulate an object into the preferred position.
The second characteristic is forceful precision gripping, in
which the cushions of the thumb and one or more of the
ngers are able to stabilize or control an object, and at the
same time tolerate large external forces, such as when
knapping a stone tool (Kivell, 2015).
In this sense, while other primates are enough skilled
to control precision grips, typically tip-to-tip or pad-to-side
grips between the thumb and index nger, these are not
generally done with strong force, and this allow the third
and uniquely human manipulative aptitude, which is the
power to squeeze gripping of cylindrical objects, that allow
ngers grip the object diagonally across the palm and the
thumb, both wrapped around the object or in line with the
forearm, for example when using a hammer (Kivell, 2015).
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
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Of course, other primates are clever enough to control
grips (using the palm) or diagonal hook grips (ngers
usually stabilized touching the palm), but neither keep
the same control that humans have to power squeeze
grip. In this sense, maybe the most critical feature to the
exclusive controlling skills of humans is our hand shape
(comparative length of the thumb and ngers), and there
is no way to forget the fth digit, that is also exceptionally
signicant during stone tool-related behaviors, because
the fth digit stabilizes the leading hand during power
squeeze grips and careful grips (e.g. during the strike
of the hammer stone), or in precision grips of the non-
dominant hand when maneuvering an object in the hand
to nd the desired position (Smouse, Focardi, Moorcroft,
Kie, Forester and Morales, 2010).
However, all these skills improved over human
development, lead to differences among cognition, as
a process of interpreting and integrating information
concerning the outside world, so it can be said that the
perceptual information and the motor commands that
represent the output of cognitive processes, are together
in order to interoperate the surroundings. More recently,
these distinctions have accepted that cognition is best
conceived as a set of processes mediating the adaptive
control of bodies in environments (Barton, 2012).
However, even if most animals are capable to
recognize, recall and using information about the places
they have been, this knowledge could potentially decrease
uncertainty about the location and accessibility of
resources, and even permit the anticipation of possibility
of danger, thanks to memory features (Dzib-Goodin,
Sanders, Yelizarov, 2017). However, there is not enough
comprehension about how animals record and use spatial
information, possibly a more realistic way or reinforced
model for animal movement would accept the possibility
of getting back to any earlier visited place even if such
locales are outside the current perception area (Smouse,
Focardi, Moorcroft, Kie, Forester and Morales, 2010).
This would mean that learning conguration of time
and space is not easy, mainly because the process
recruit many systems in order to manage data-acquisition
mechanisms to produce a specic output, since all that
requires much memory and computation of specic
information (Dzib-Goodin, Sanders, Yelizarov, 2017;
Forterre, Gribaldo, Brochier, 2005).
So, it is important to realize that both cognitive
and motor functions involve the learning of sequential
actions. These sequences are adjusted with control by
particular arrangements mediated by both executive
function and automaticity, because learning complex
sequences involves effective performance of executive
processes, this have been demonstrated an overlap in
the supplementary motor cortex and other brain regions,
such as the cerebellum, basal ganglia premotor cortex,
thalamus, ventrolateral premotor cortex, and precuneus,
with increased activations at increased levels of complexity
(Leisman, Moustafa and Shar, 2016).
Other special feature associated with movement
was the shift of body-space structure associated with
the appearance of hominin bipedalism (Dzib-Goodin,
Yelizarov, 2016), this might have another effect to
development specic brain areas, and specically
extended to opercular cortex. Such neural development
(construction of a neural niche) could enable the managing
of abstract information, separate from actual physical
limitation, by applying and re-using existing codes for
spatial information processing to understand novel mental
purposes (construction of the cognitive niche), and this
could give as a result the development of language. This
is believed because focused manipulation of the body
image in space, demanded for tool use, would have
rushed collaborating relations between the neural and
cognitive niches, and because tool use needs a change
of numerous bodily and spatial skills as well as logical
and sequential relations of action components (Iriki and
Taoka, 2012).
In this sense, tools engage cognitive brain functions,
linked with ne movements, including language (Dzib-
Goodin, Sanders, Yelizarov, 2017), since they were
created one after another and shared into hominid
environments as essential elements, that created what is
known as construction of the ecological niche. Sooner or
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
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later, a human-modied environment puts emphasis on
following generations to familiarize to it, conceivably by
getting additional resources for other relevant tissues.
Epigenetically this encouraged plasticity (including
developmental or learning processes) so would contribute
to allow that extra genomic information could be spread
among generations via mutual exchanges between
ecological, neural and cognitive domains of niches, which
may have funded to hominid evolutionary processes. This
scenario would nd the human brain as part of an evolving
holistic ecosystem (Iriki and Taoka, 2012; Godfrey-Smith,
Movement process and adaptation to the environment
Once tools and language began to interact to
create better environments, were added more cognitive
processes. It is accepted that after modern humans
departed from sub-Saharan Africa, over 50 000–100 000
years ago, physical changes were necessary to diverse
environments. In this regard, it is thought that when human
populations were exposed to additional environmental
changes, this produced cultural innovations, such as the
increase of agriculture, which gave rise to new selective
compressions linked to pathogen exposures and dietary
changes and this at the same time, altered the frequency
of individual adaptive alleles, so it is easy to believe that
natural selection also make up the overall genetic and
brain architecture of adaptive traits (Olson, Knoester,
Adami, 2016).
From this perspective, other process was important,
specically animal-grouping behavior, which had
important consequences for social intelligence and
collective cognition, since grouping behaviors are
persistent across all forms of life. As an example, is
possible to mention swarming as a grouping behavior,
where animals synchronize their movement with the rest
of their group to maintain a cohesive unit. In this sense,
swarming may increase matting success, spread foraging
efciency, or enable the group to resolve problems that
would be impossible to solve alone, plus there is indication
of cerebellar development and participation in different
cognitive functions, depending the kind of grouping
behavior, suggesting a link between neocortex size and
social group size (Barton, 2012).
Also, swarming behaviors could protect group
members from predators in several ways, and in this
regard, swarming can improve group vigilance, reduce the
chance of being encountered by predators, and reduce
individual possibility of being attacked, allowing an active
protection against predators, or reduce reducing predator
assault efciency by confusing the predator (Olson,
Knoester, Adami, 2016).
In other words, it is important to move efciently into
the physical space, alone or in group, in order to get a
better opportunity to survive, and by this reason Darwin
(cited by Kivell, 2015), rst suggested that the introduction
of bipedalism was directly connected to tool use as a way
to free the hands and expand the locomotion. So, the
relationship regarding motor function and cognition can
be understood, in part, in the context of the evolution of
human bipedalism, which helped as a signicant basis for
the evolution of the human neocortex as it is among the
most complex and sophisticated of all movements (Jeong
and Di Rienzo, 2014).
This gave humans a unique capacity to relate
gravitational forces as a direct result of the existence of
the erect position. The basis of the continuation of this
genetic mutation is based on the notion that bipedalism
had created larger pools of neurons. It is debated that the
same evolutionary process has permitted to develop the
binding of the motor system into synchronous, rhythmic,
purposeful movement, which expanded to eventually allow
for cognitive binding and perception (Leisman, Moustafa
and Shar, 2016).
This required a change for the hips and pelvis,
not only in motion but muscle innervations (Dunbar,
Horak, Macpherson, Rushmer, 1986), since walking
and running implicate more support from joints and
ligaments (Muehelenbein, 2015) and changes in knees as
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
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a mechanical motor to use and produce energy to move
erected (Hogervost, Bourna, & de Vos, 2009). So this
adaptation provided a better synchronization and efciency
to hip extensor mechanism that create a different system
comparted with other hominids (McHenry, 1975), that
pushed homos from the arboreal walking to the ground
(Hanna, Grabatosky, Rana and Schmitt, 2017).
This development allows humans to move further,
and explore other ways to survive, like creating harborage,
nd different kind of food and be part of other groups.
While memory of food locations and higher cognition
may limit the benets of random walk schemes (Dzib-
Goodin, Sanders, Yelizarov, 2017), so called Lévy walks
may have result from with the evolution of a hunting and
gathering lifestyle in human lineages. Lévy walks are an
unsystematic walk creating a new strategy used by a wide
variety of organisms when searching for food (Raichlen,
Wood, Gordon, Mabulla, Marlowe and Pontzer, 2014).
This kind of search implicates frequently short move
steps (dened as the distance traveled before pausing
or changing direction) merged with unusual longer move
steps (Smouse, Focardi, Moorcroft, Kie, Forester and
Morales, 2010).
This movement arrangement can be essential to
understand how humans perceive and interrelate with the
world within a wide range of ecological frameworks, and
it might be adaptive behavior to solve food distribution
arrangements on the landscape. The widespread use of
this movement pattern among species with great cognitive
disparity insinuates a link between hunting patterns within
different organisms, including humans (Raichlen, Wood,
Gordon, Mabulla, Marlowe and Pontzer, 2014).
As a result of the interaction with environment, larger
regions of posterior parietal cortex and frontal motor cortex
become part of special networks dedicated to generating
different series of movements, consequently, motor
areas include primary motor cortex, ventral (PMv), and
dorsal (PMd) premotor cortex, the supplementary motor
area (SMA), and the frontal eye eld (FEF). However,
movement cannot do too much without the interaction
with senses, so somatosensory regions incorporate the
four areas of anterior parietal cortex. As a result, primary
motor cortex and dorsal and ventral premotor areas are
well-known as cortical areas, and each of these areas has
a somatotopic representation of minor activities of body
parts (Kass, 2008; Kass, 2012b). Curiously, these areas
are compromise in movement disorders such as apraxia
(Murillo Duran, 2007).
This paper is just a brief and not exhaustive view
of movement process as a key of evolution of species
and human cognition, specically from prokaryote to
eukaryote cells to human cognition. Millions of years have
been needed to draft more than one biology model of our
From this perspective, movement process is not only
important in large scale of the universe, since it keeps
galaxies and planets in a perfect dance, but it has an
impact into cells, in order to create a diversication of
functions, adaptation and physical features.
One scenario explored is that phagocytosis could
be a key to change the evolutionary rhythm of life, and
actin proteins created new options to motility, that is why a
globular major component of the cellular cytoskeleton and
one of the most abundant cellular proteins.
However, It was needed still a long period of time
before see a primitive nervous system, probably because
the advance of the sensory processes, that beside
motor behavior began to create the neuronal networks in
the rst nervous systems that is possible to appreciate
among different species. As a result the human brain with
a sensory motor system capable not only to understand
the environment, but also manipulate its own resources to
create adaptive answers to the environment.
Once that human brain was capable to recognize
itself is physical space and time, walking create a cultural
revolution allowing even more connections, and allowing
memory to create marks to recognize the environment.
Some believe thanks to the use of tools, communication
began in other ways more than just calls, and this create
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
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a cognitive niche to connect with the rest of the human.
We have explored in other articles than memory
was result of movement, so of course explains why is
so important to learning process. From psychological
standpoint, several authors have claimed that movements
seen as physical activity are important to learning process,
but in our perspective, they are not capable to explain why
this relationship is so important to human brains.
That is why this complex process must be seen
from different perspectives, from microbiology, genetic,
evolution, cultural, cognitive, clinic and even artistic point
of view, and certainly each area has many more to say,
because it is, from our perspective, very important to
understand how cognition built human brains, that is just
one example of evolution of species.
We deeply believe human brain is not the last draft
of evolution, cognitive processes have been modulated
based environmental needs and those changes that prove
to be important over the population will become part of the
repertory and structures of the brains. This is not a human
design, but a species mechanism to survive.
Conict-of-Interest Statement: authors declare that
no competing or conict of interest exists. Received: 03/06/2017
Accepted: 10/07/2018
Evolution of movement / Alma Dzib-Goodin; Daniel Yelizarov
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From an evolutionary point of view, the process of memory evolved specially among mammals as a cognitive skill needed to conceive the future. This was possible using known elements in order to respond to the environment, however the process of memory cannot work independent of other systems such as learning, sleeping, movement and attention because memories are created first for self-preservation and second, as a tool for survival of the species. The evolution of the memory process requires the recruitment of different brain structures that primarily support the hippocampus, but additionally, involves some molecular adaptations such as the role of glucocorticoids, proteins and kinase family participation, which demonstrates a flexible activity that adapts and responds to environmental needs. The epigenetic processes possible thanks the modification of core histones, has an important role to regulate these processes through cellular signals that integrate molecular information in transcription and transduction, modulating synaptic plasticity under the restriction of calcium channels.
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Active matter systems, and in particular the cell cytoskeleton, exhibit complex mechanochemical dynamics that are still not well understood. While prior computational models of cytoskeletal dynamics have lead to many conceptual insights, an important niche still needs to be filled with a high-resolution structural modeling framework, which includes a minimally-complete set of cytoskeletal chemistries, stochastically treats reaction and diffusion processes in three spatial dimensions, accurately and efficiently describes mechanical deformations of the filamentous network under stresses generated by molecular motors, and deeply couples mechanics and chemistry at high spatial resolution. To address this need, we propose a novel reactive coarse-grained force field, as well as a publicly available software package, named the Mechanochemical Dynamics of Active Networks (MEDYAN), for simulating active network evolution and dynamics (available at This model can be used to study the non-linear, far from equilibrium processes in active matter systems, in particular, comprised of interacting semi-flexible polymers embedded in a solution with complex reaction-diffusion processes. In this work, we applied MEDYAN to investigate a contractile actomyosin network consisting of actin filaments, alpha-actinin cross-linking proteins, and non-muscle myosin IIA mini-filaments. We found that these systems undergo a switch-like transition in simulations from a random network to ordered, bundled structures when cross-linker concentration is increased above a threshold value, inducing contraction driven by myosin II mini-filaments. Our simulations also show how myosin II mini-filaments, in tandem with cross-linkers, can produce a range of actin filament polarity distributions and alignment, which is crucially dependent on the rate of actin filament turnover and the actin filament's resulting super-diffusive behavior in the actomyosin-cross-linker system. We discuss the biological implications of these findings for the arc formation in lamellipodium-to-lamellum architectural remodeling. Lastly, our simulations produce force-dependent accumulation of myosin II, which is thought to be responsible for their mechanosensation ability, also spontaneously generating myosin II concentration gradients in the solution phase of the simulation volume.
Vertical climbing is an essential behavior for arboreal animals, yet limb mechanics during climbing are poorly understood and rarely compared to those observed during horizontal walking. Primates commonly engage in both arboreal walking and vertical climbing, and this makes them an ideal taxa in which to compare these locomotor forms. Additionally, primates exhibit unusual limb mechanics compared to most other quadrupeds, with weight distribution biased towards the hindlimbs, a pattern that is argued to have evolved in response to the challenges of arboreal walking. Here we test an alternative hypothesis that functional differentiation between the limbs evolved initially as a response to climbing. Eight primate species were recorded locomoting on instrumented vertical and horizontal simulated arboreal runways. Forces along the axis of, and normal to, the support were recorded. During walking, all primates displayed forelimbs that were net braking, and hindlimbs that were net propulsive. In contrast, both limbs served a propulsive role during climbing. In all species, except the lorisids, the hindlimb produced greater propulsive forces than the forelimb during climbing. During climbing, the hindlimb tends to support compressive loads, while the forelimb forces tend to be primarily tensile. This functional disparity appears to be body-size dependent. The tensile loading of the forelimb versus the compressive loading of the hindlimb observed during climbing may have important evolutionary implications for primates, and it may be the case that hindlimb-biased weight support exhibited during quadrupedal walking in primates may be derived from their basal condition of climbing thin branches.
The purpose of this work is to analyze paraxial difficulties i.e, functional disorders in movement resulting from cerebral tissue lesions. In accordance with the literature on the subject, the following definition, the following definition of apraxia has been accepted: "Apraxia is inability in properly executing kinetic tasks without impairment or loss of motor or sensory functions or ataxia with would condition such inability". "Proper execution" used in this definition concerns not only the effect of the action but also means of its realization. "Kinetic tasks" signify all aspects of motor activity defined by instructions, irrespective of the type of instruction (verbal or gestures), and regardless of whether it required -in the final effect- imitating the movement of the investigator or whether they were performed independently. The methodology has been based in the principle on Luria’s works. Deliberations on praxial disorders were based on investigation results embracing 90 patients with focal cerebral lesions. In fifty cases, changes were localized in the left cerebral hemisphere, in forty cases -in the right hemisphere. The summing up the results concerning a global comparison between cerebral hemispheres, indicate the following regularities: Results achieved made it possible to form the opinion that not all of the generally accepted tests investigating praxia in persons with cerebral lesions are solved faultlessly by healthy individuals; thus, a faulty execution should not always be regarded as a sign of pathological functioning of the cerebral tissue as a result of lesion.
Self-assembly of two important components of the cytoskeleton of eukaryotic cells, actin microfilaments and microtubules (MTs) results in polar filaments of one chirality. As is true for bacterial flagella, in actin microfilaments, screw direction is important for assembly processes and motility. For MTs, polar orientation within the cell is paramount. The alignment of these elements in the cell cytoplasm gives rise to emergent properties, including the potential for cell differentiation and specialization. Complex MTs with a characteristic chirality are found in basal bodies and centrioles; this chirality is preserved in cilia. In motile cilia, it is reflected in the direction of the effective stroke. The positioning of the basal body or cilia on the cell surface depends on polarity proteins. In evolution, survival depends on global polarity information relayed to the cell in part by orientation of the MT and actin filament cytoskeletons and the chirality of the basal body to determine left and right coordinates within a defined anterior–posterior cell and tissue axis. This article is part of the themed issue ‘Provocative questions in left–right asymmetry’.
Axonal growth is a complex phenomenon in which many intra- and extra-cellular signals collaborate simultaneously. Two different compartments can be identified in the growing axon: the growth cone, the leading tip that guides and steers the axon, and the axonal shaft, connecting the soma to the growth cone. The complex relations between both compartments and how their interaction leads the axon to its final synaptic target remain a topic of intense scrutiny. Here, we present a continuum and computational model for the development of the axonal shaft. Two different regions are considered: the axoplasm, filled with microtubules, and the surrounding cortical membrane, consisting mainly of F-actin, Myosin II motor proteins and the membrane. Based on the theory of morphoelasticity, the deformation gradient is decomposed into anelastic and viscoelastic parts. The former corresponds to either a growth tensor for the axoplasm, or a composition of growth and contractile tensors for the cortical membrane. The biophysical evolution for the anelastic parts is obtained at the constitutive level, in which the polymerization and depolymerization of microtubules and F-actin drive the growth, while the contractility is due to the pulling exerted by the Myosin II on the F-actin and depends on the stress. The coupling between cytoskeletal dynamics and mechanics is naturally derived from the equilibrium equations. The framework is exploited in two representative scenarios in which an external force is applied to the axonal shaft either along the axis or off the axis. In the first case three states are found: growth, collapse and stall. In the second case, axonal turning is observed. This framework is suitable to investigate the complex relationship between the local mechanical state, the cytoskeletal polymerization/depolymerization rates, and the contractility of the cortical membrane in axonal guidance.
An analysis of the genes that make up the DRβ region will extend our information about the complexity of the HLA class II system, since the serologically detectable polymorphism is located mainly on the DRβ chain. Probably this analysis will be most informative if we look at the 5′ ends of the genes coding for the variable stretches of a DRβ chain [1].