THE BRAIN IN THE

VERTEBRATE SERIES
AN IN-DEPTH EXPLORATION FROM CYCLOSTOMES TO MAMMALS

David López Torres

Comparative anatomy of vertebrates
 Table of contents

Cyclostomes .................................................................................................................................. 2
Transition to fish ............................................................................................................................ 3
Amphibians: The bridge to terrestrial life ..................................................................................... 4
Reptiles: The rise of cognitive complexity ..................................................................................... 5
Birds: The avian enigma ................................................................................................................ 6
Mammals: Pinnacle of vertebrate brain evolution........................................................................ 7
HOX genes: Orchestrating vertebrate brain evolution .................................................................. 8
Experiments and Anatomical Discoveries: Unravelling the Mysteries of Vertebrate Brains ......... 9
Conclusion ................................................................................................................................... 10
Bibliography................................................................................................................................. 11

Introduction
Starting from the primitive cyclostomes and advancing to the sophisticated mammals, each stage
in this evolutionary development has played a role in the remarkable diversity observed in the
anatomy and configuration of vertebrate brains. This essay seeks to examine the intricacies of
the vertebrate brain throughout its series, encompassing cyclostomes, fish, amphibians, reptiles,
birds, and mammals. Additionally, it will investigate the influence of HOX genes, pivotal
experiments, and anatomical revelations that have contributed to shaping our comprehension
of the evolution of vertebrate brains.

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 1. Cyclostomes

Cyclostomes, exemplified by lampreys and hagfish, provide insight into the beginnings
of vertebrate brains. Their brains exhibit a basic structure, comprising a small spinal cord
and a primitive brain region referred to as the rhombencephalon. It is significant to note
the lack of features such as the cerebellum and neocortex (present in more advance
vertebrates), suggesting a less complex neural organization.

In cyclostomes, the brain is differentiated into forebrain, midbrain, and hindbrain

regions, though not as prominently as in more evolved vertebrates, indicating an early
evolutionary phase. Despite simplicity, the cyclostome brain demonstrates adaptability,
enabling success in diverse environments. This adaptability is evident not only in basic
brain organization but also in the development of specialized structures like the olfactory
and pineal organs, showcasing the versatility of brain structures across different
vertebrate lineages:

• Olfactory system: Due to their dependence on the sense of smell, lampreys possess
a highly developed olfactory bulb and related structures. This adaptation highlights
the significance of olfaction in their sensory capabilities. Additionally, their well-
developed olfactory system is essential for tasks such as locating prey and navigating
their environment.

• Pineal organ: Cyclostomes have a pineal organ that plays a role in regulating
biological rhythms and responding to light cues in the environment. In lampreys, the
pineal organ is particularly acknowledged as a vital photoreceptive structure. This
organ not only helps coordinate biological rhythms but also assists in sensing
alterations in light conditions.

These insights connect to the broader narrative of fish brain evolution, portraying
ancestral conditions pivotal for the emergence of advanced brain structures in higher
vertebrates.

Illustration 1: Central nervous system of a lamprey: forebrain (blue), midbrain (green), hindbrain (orange), choroidal
tissue (yellow) and spinal cord (pink).

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 2. Transition to fish

The transition from cyclostomes to fish marks a significant step in vertebrate brain
evolution. Fish, as the first jawed vertebrates, exhibit a more developed brain structure
compared to cyclostomes. The brain is divided into three primary regions: the forebrain,
midbrain, and hindbrain.

One notable advancement is the emergence of the cerebellum, responsible for motor
coordination and balance. This development reflects the adaptive advantage conferred
by enhanced motor control in aquatic environments. The forebrain also undergoes
considerable expansion, setting the stage for increased sensory processing and cognitive
abilities in subsequent vertebrate groups. In the case of teleost fishes, a significant
evolution of the brain occurred, particularly an enlargement of the forebrain,
encompassing the olfactory bulbs and telencephalon enhancing sensory processing and
the development of complex behaviours.

Fish brains exhibit a clear correlation between brain structure and ecological demands,
underscoring the role of natural selection in shaping vertebrate neuroanatomy. As fish
adapted to various ecological niches, specific brain regions became specialized for
distinct functions. For instance, species inhabiting environments with intricate spatial
navigation requirements might possess an enlarged cerebellum. These adaptations
encompass well-developed olfactory bulbs for odour detection, advanced vision centres,
and lateral line systems for perceiving water movement and pressure changes. We will
find alterations in brain size, sensory structures, and cognitive abilities to navigate
specific challenges.

With the study of fish brains, we have taken another step in the evolution of vertebrates.
Considering that these constitute one of the oldest groups, this allows us to understand
and compare, with our next group, the amphibians, characteristics that differ and mark
an evolution in the brain.

Illustration 2: Dorsal view of the brain of a rainbow trout (Oncorhynchus mykiss)

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 3. Amphibians: The bridge to terrestrial life

Amphibians serve as crucial intermediaries bridging the realms of aquatic and terrestrial
vertebrates, undergoing significant brain adaptations in their transitional journey. Their
brains, marked by an expanded forebrain supporting sensory perception and the
emergence of the limbic system for emotional responses, showcase sophistication
tailored to diverse habitats.

This evolutionary narrative unfolds in frogs, toads, salamanders, and newts, constituting
a captivating chapter in vertebrate evolutionary history. As a transitional group linking
fish and reptiles/birds, amphibians undergo brain adjustments to navigate life both in
water and on land.

Examining the brain divisions into forebrain, midbrain, and hindbrain, intriguing facts
emerge:

• Development of Olfactory Bulbs: Amphibians, reliant on their sense of smell, boast

well-developed olfactory bulbs.

• Optic Tectum and Vision: The remarkably large optic tectum in amphibians,
particularly aquatic species, underscores the paramount role of vision.

• Cerebellum and Motor Control: The amphibian cerebellum, crucial for motor control,
adapts to facilitate refined locomotion upon the transition to land.

The evolution of amphibian brains not only illuminates broader trends in vertebrate
brain development during the transition from aquatic to terrestrial environments but
also provides crucial insights into the origins of sophisticated brain structures observed
in mammals and birds. This journey through amphibian brain evolution showcases the
intricate interplay between structure, function, and environmental adaptation,
contributing to our understanding of vertebrate neuroanatomy.

Illustration 3: central nervous system of an amphibian

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 4. Reptiles: The rise of cognitive complexity

Reptiles occupy a pivotal position in the evolutionary trajectory of vertebrates, marking

a significant advancement in both brain size and complexity. Among the distinctive
features characterizing reptilian brains, the development of the cerebral cortex stands
out as a key contributor to higher cognitive functions.

Structural modifications in the reptilian brain encompass the division of the cerebrum
into discernible lobes, a structural enhancement fostering heightened sensory
processing and improved motor coordination. Noteworthy is the ongoing evolution of
the olfactory system, underscoring the paramount role of chemical communication in
guiding reptilian behaviour.

The evolutionary triumph of reptiles is further underscored by the emergence of the

limbic system, a critical development indicating enhanced emotional processing and
advanced learning capabilities. These adaptations lay the groundwork for the
subsequent evolution of even more intricate brain structures in avian and mammalian
lineages.

The reptilian brain's capacity for intricate cognitive functions and its adaptability to
environmental challenges highlight the success of reptiles in diverse ecosystems. Their
brain's structural refinements not only serve as a testament to the efficiency of reptilian
neuroanatomy but also provide valuable insights into the evolutionary processes
shaping the development of sophisticated brain structures observed in more advanced
species such as birds and mammals. The reptilian brain serves as a crucial link in
understanding the continuum of vertebrate brain evolution, showcasing the dynamic
interplay between structure, function, and environmental adaptation.

Illustration 4:Scheme of the brain of an alligator

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 5. Birds: The avian enigma

The evolution of the avian brain is a captivating subject within vertebrate biology. Birds,
descendants of theropod dinosaurs, have developed sophisticated brains with a high
degree of encephalisation enabling complex behaviours such as problem-solving, vocal
learning, and intricate social interactions. Key features contribute to their brain
evolution:

o Forebrain Development: The well-developed forebrain, particularly the

telencephalon, enhances cognitive functions, facilitating learning and complex tasks.
Species like parrots and songbirds showcase complex vocal learning associated with
specific neural circuits.

o Brain-to-Body Mass Ratio: Intelligent birds often have a high brain-to-body mass
ratio, indicative of advanced cognitive capabilities.

o Social Complexity: Birds in socially complex environments, like crows, exhibit more
developed brains, aiding in advanced problem-solving.

The avian brain's evolution is shaped by ecological and behavioural factors, offering
insights into convergent evolution of intelligence. Avian brains, characterized by
encephalization, the pallium (Equivalent to the neocortex in mammals, it is associated
with the processing of complex information and intelligent behaviour), and specialized
neural circuits, represent a remarkable convergence of traits associated with advanced
cognitive abilities.

Illustration 5: Scheme of a goose brain

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 6. Mammals: Pinnacle of vertebrate brain evolution

Early mammals exhibited modest brain sizes and likely engaged in uncomplicated
behaviours. The shift from reptiles to mammals approximately 200 million years ago
signified the commencement of mammalian brain evolution, paving the way for
increased complexity in cognitive functions and behaviours.

This group is the pinnacle of vertebrate brain evolution. The brains of mammals began
to increase in size, and with this, there is an observed trend that, the larger the size, the
greater the complexity. Larger brains are usually associated with more complex and
improved cognitive abilities, such as problem-solving, sensory perception, motor
commands, spatial reasoning, and language. This allowed mammals to adapt to a greater
variety of ecological niches, positioning them as the predominant group in ecosystems,
this leads us to have a higher encephalization quotient (a measure of brain size relative
to body size), especially prominent in primates. This highlights the significance of the
brain in controlling complex behaviours.

The mammalian brain is divided into several distinct regions, including the cerebral
cortex, limbic system, and cerebellum. The cerebral cortex, unique to mammals,
undergoes significant expansion, contributing to the complexity of sensory processing
and higher-order cognition.

The limbic system, particularly the hippocampus and amygdala, plays a significant role
in emotional regulation and memory formation. This development correlates with the
intricate social behaviours observed in mammals, emphasizing the role of the brain in
shaping complex social structures.

Illustration 6: Scheme of different mammal’s brain

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 7. HOX genes: Orchestrating vertebrate brain evolution

HOX genes are a conserved family crucial for embryonic development, regulating spatial
and temporal gene expression, wield considerable influence over the shaping of the
vertebrate brain. These genes oversee the spatial and temporal expression that
determines segment identity along the anterior-posterior axis of the developing embryo.
In the context of the brain, HOX genes play a pivotal role in regionalization and
specialization, with experimental studies providing insights into the molecular
mechanisms that underlie vertebrate brain evolution.

While the primary function of HOX genes is in the body plan and segment identity during
embryonic development, they also influence the patterning and development of the
vertebrate brain.

In the developing vertebrate hindbrain, encompassing structures like the medulla and
pons, HOX genes demonstrate segmental expression known as rhombomeric
segmentation, this segmentation is vital for hindbrain development, contributing to the
identity of each rhombomere and forming distinct hindbrain structures.

This genes also plays a role in cranial nerve development, impacting neurons associated
with these nerves and governing various head and neck functions. While their primary
influence is in the hindbrain, HOX genes also affect the forebrain and midbrain,
particularly during early embryonic development.

The HOX genes are conserved throughout evolution, despite variations in the cerebral
complexity among different species. These genes help us understand the formation of
different brain regions, and their regulation is essential for establishing the complex
structure and function of the vertebrate brain.

Illustration 7:Position of the Hox genes along the chromosome and the corresponding area of the body of Drosophila
melanogaster (subject in which this type of genes was discovered) where they are expressed.

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 8. Experiments and anatomical discoveries: Unravelling the mysteries of
vertebrate brains

Below are described the most notable experiments in the field of neuroscience:

o Split-Brain Studies by Roger Sperry (1960s): The term "split-brain" refers to a surgical
procedure called callosotomy, which involves cutting the corpus callosum and the
bundle of nerve fibers connecting the two hemispheres. This procedure was
primarily performed to alleviate severe cases of epilepsy, where seizures originated
in one hemisphere and spread to the other through the corpus callosum.
With these experiments, they discovered the independence of the hemispheres
(both brain hemispheres could operate independently), the lateralization and
specialization of function (with behaviours and thoughts associated with specific
hemispheres), and the challenges in communication (by cutting the corpus callosum,
each part of the brain became "independent," rendering patients unable to verbalize
an act that had been processed by the opposite hemisphere).

o Mirror Neurons (1990s): The discovery of mirror neurons was carried out by a team
led by the Italian Giacomo Rizzolatti. These are a type of brain cell that activates
when an individual performs an action, just as when they observe someone else
performing the same action, suggesting a neural reflection.
This discovery was made while, originally, were studying motor neurons in the brains
of macaques. It is believed that the function of mirror neurons plays a crucial role in
the imitation and empathy of other actions and is considered a fundamental
mechanism for social interaction. Subsequent research suggests that mirror neurons
also exist in humans (such as the act of yawning). Neuroimaging studies using
techniques like functional magnetic resonance imaging (fMRI) have identified brain
regions in humans that show similar patterns of activation during the execution and
observation of actions.

o Human Connectome Project, HCP (2009): This project is a scientific collaboration

initiative with the primary goal of comprehensively mapping the functional and
structural connectivity of the human brain. This involves understanding the different
parts of the brain and how they are interconnected using techniques such as
structural magnetic resonance imaging (MRI), functional magnetic resonance
imaging (fMRI), and diffusion tensor imaging (DTI). The study initially focused on
twin pairs to explore genetic variability in brain connectivity but also included single
individuals, and they share all the information, making it publicly accessible.
The data collected by the HCP has led to significant advances in understanding brain
architecture and has enabled research on individual variability in connectivity, the
relationship between brain structure and function, as well as the connection
between brain connectivity and various neuropsychiatric disorders.

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 9. Conclusion

The evolutionary journey across vertebrates unfolds a compelling narrative of innovation

and adaptation. From the rudimentary brain of cyclostomes to the sophisticated brains
of mammals, each stage reflects the impact of selective pressures and ecological
challenges on the vertebrate nervous system.

The significance of HOX genes in coordinating developmental processes shaping brain

evolution highlights the intricate molecular dynamics at play. Through experiments and
anatomical revelations, our comprehension of vertebrate brains has significantly
advanced, providing unparalleled precision.

As we get deeper into the intricacies of the vertebrate brain, the collaboration between
neurobiology, genetics, and palaeontology promises ongoing revelations. The ongoing
saga of vertebrate brain evolution invites scientists to explore the captivating crossroads
of genetics, anatomy, and behaviour with increasing depth.

Illustration 8: Comparative scheme of the different brains, example of different orders in the vertebrate subphylum

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