Developmental Zoology

Comparison of Gametogenesis Across Animal Classes: Insights from

Chicken, Frog, Human, and Starfish
Jemimah A. Cadag1, Christian C. Contacto1, Catherine Kaye R. Ocfemia1,
Kurt John B. Palivino1, Patricia E. Peran1
1
Department of Biology, College of Science, Bicol University
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ABSTRACT

Gametogenesis is the process by which life can continue to proliferate and exist, it occurs for the
purpose of sexual reproduction and is an important part of individual maturation often signaling the
transition from childhood to adulthood. Primordial germ cells in the gonads give rise to primary
gametocytes, these contain two copies of each chromosome being known as diploid cells. Now the
process by which gametogenesis occurs and its mechanisms are different for each species due to
differences in morphology, environmental conditions and specializations that may have come about from
evolution. Understanding the nuances of gametogenesis not only illuminates the marvel of reproductive
biology but also unveils evolutionary adaptations and reproductive strategies tailored to diverse ecological
niches. This paper embarks on a comparative review of gametogenesis across various animal classes and
their representative species, ranging from chickens and frogs to humans and starfish. Through
comparative analysis, intriguing variations in gametogenesis among species are revealed, influenced by
ecological, physiological, and evolutionary factors. This highlights the fundamental aspect of sexual
reproduction, showcasing both universal principles and species-specific adaptations. By unraveling the
complex interplay between genetic, developmental, and environmental factors shaping reproductive
strategies.

Keywords: Gametogenesis, Comparative, Representative, Morphology , Evolutionary

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INTRODUCTION ovaries. Both processes culminate in the

Gametogenesis, the intricate process
through which gametes develop, serves as a
cornerstone of sexual reproduction across the
animal kingdom. Spermatogenesis in males and
oogenesis in females orchestrate this vital
process, with each occurring within specialized
reproductive organs. Spermatogenesis takes
place within the seminiferous tubules of the
testes, while oogenesis unfolds within the
production of haploid cells primed for
fertilization.
Spermatogenesis commences with
diploid spermatogonia undergoing mitotic
divisions to generate primary spermatocytes.
These primary spermatocytes then undergo
meiosis, resulting in the formation of haploid
spermatids, which subsequently mature into
spermatozoa. Conversely, oogenesis involves the
Developmental Zoology
growth and maturation of primary oocytes into
secondary oocytes, accompanied by the
formation of polar bodies through meiotic
divisions. This process ultimately yields a single
haploid ovum and two or three polar bodies,
which are subsequently discarded.
Understanding the nuances of
gametogenesis not only illuminates the marvel
of reproductive biology but also unveils
evolutionary adaptations and reproductive
strategies tailored to diverse ecological niches.
This paper embarks on a comparative review of
gametogenesis across various animal classes and
their representative species, ranging from
chickens and frogs to humans and starfish. By
exploring these differences, we gain insights into
the remarkable diversity and intricacies of
reproductive mechanisms, shedding light on
fundamental principles underlying sexual
reproduction.
Through comparative analysis,
intriguing variations in gametogenesis among
species are revealed, influenced by ecological,
physiological, and evolutionary factors. This
highlights the fundamental aspect of sexual
reproduction, showcasing both universal
principles and species-specific adaptations. By
unraveling the complex interplay between
genetic, developmental, and environmental
factors shaping reproductive strategies, this
study contributes to our understanding of
reproductive biology.
Male and female humans exhibit
internal sexual dimorphism, characterized by
distinct anatomical and physiological differences
encompassing unique reproductive organs and
hormonal attributes. In humans, males produce
motile gametes called sperm, in contrast to the
non-motile eggs produced by females. While
some other species may follow a similar pattern,
others, such as frogs and starfish, are influenced
by seasonal events. This is also applicable to
post-gametogenesis developmental processes
(Gilberts, 2000; Wessel et al., 2010; Mariante et
al., 2010).
The production of human reproductive
cells occurs in reproductive organs. In males,
sperm production, release, and storage are
facilitated by the testes, tubular networks, and
epididymis, respectively. Moreover, ovaries are
essential for egg production, and the fallopian
tubes transport them to the uterus for
fertilization to occur. Some species exhibit
commonalities in the basic functions of the
reproductive organs concerning their gametes.
However, specific structures are modified
depending on environmental conditions and
reproductive strategies.
METHODOLOGY
Literature review and comparative
analysis were conducted to examine the process
of gametogenesis in chickens, frogs, humans,
and starfish. Relevant scientific articles,
textbooks, and online databases were consulted
to gather information on the anatomy,
physiology, and molecular mechanisms
underlying gametogenesis in each species.
DISCUSSION
In humans and other mammals, the
onset of puberty marks the beginning of the
intricate hormonal interplay that governs
gametogenesis. Gonadotropin-releasing
hormones (GnRH) from the hypothalamus
kickstart the release of follicle-stimulating
hormone (FSH) and luteinizing hormone (LH)
Developmental Zoology
from the pituitary gland. These hormones
orchestrate the production of testosterone in
males, essential for spermatogenesis, and
regulate the menstrual cycle and ovulation in
females. Additionally, inhibin plays a crucial
role in modulating spermatogenesis by inhibiting
GnRH and FSH release once sperm count levels
are sufficient. Sertoli cells initiate the production
of Inhibin to slow down spermatogenesis
through the inhibition of GnRH and FSH.
Environmental factors, such as
temperature and psychological stress, can
significantly influence gamete production and
quality. Sperm viability, for instance, is affected
by temperature, with optimal motility observed
at lower temperatures. Psychological imbalance
can disrupt hormonal harmony, impacting
reproductive cycles.Some organisms including
humans are influenced by environmental factors
that greatly affect the production and quality of
their gametes. Temperatures are a highly
significant factor to consider for sperm viability.
From a study of (Appell et al,1977) sperm
motility is best preserved at 20ºC. At higher
temperature (37ºC), motile sperms are reduced
in count.
Another factor to consider is the
psychological imbalance that disrupts the
harmony in reproductive hormones. While
experiencing depressive episodes due to
hormonal imbalance during specific stages of
reproductive cycles is common, it's important to
note that the imbalance can also be a
consequence of such episodes. Although a study
regarding the fertility success among women
undergoing in vitro fertilization claims that there
is no significant relationship between stress
affecting pregnancy or delivery (Klonoff-Cohen
et al., 2001). The same study also claims that
only biologic end points are considered
significantly affected by stress manifested by
low retrieval rate and of fertile oocytes . It is
important to note however, that counseling
alleviates psychological distress which
significantly increases pregnancy rate (Rooney
& Domar, 2018).
Gametogenesis displays remarkable
diversity across species. In humans, the
development of germ cells leads to the formation
of sperm in males and eggs in females. The
development of germ cells begins with
primordial germ cells (PGCs), The specification
of PGCs in mammals is influenced by Bone
Morphogenetic Protein (BMP) signals from the
extraembryonic ectoderm and visceral endoderm
(de Sousa Lopes et al., 2004; Lawson et al.,
1999). With BMPs influencing the PGCs, in
human spermatogenesis, spermatogonia undergo
a series of cell divisions to form spermatocytes.
Similarly, in females, oogenesis occurs through
the mitosis of oogonia, forming the primary
oocytes. From the primary oocyte, the formation
of the secondary oocyte occurs, followed by
further division, resulting in the development of
a mature ovum. What differentiates mammalian
gametogenesis from others is its asymmetrical
division in oogenesis, influenced by the position
of the cytoskeleton during meiosis I and meiosis
II (Brunet, S. & Verlhac, M. H., 2010). This
non-universal feature highlights the
evolutionarily driven gametogenesis strategies
adapted by a species.
The process by which gametogenesis
occurs and its mechanisms are different for each
species due to differences in morphology,
environmental conditions and specializations
that may have come about from evolution. A
good comparison of such a case is that of the sea
star gametogenesis and human gametogenesis.
In starfish, both males and females produce large
numbers of gametes (sperm and eggs)
simultaneously while in humans, males produce
millions of sperm daily, while females are born
with a fixed number of eggs and release one egg
per menstrual cycle. Starfish gametogenesis
occurs in specialized structures called gonads,
Developmental Zoology
which are located in the arms. Human
gametogenesis occurs in the gonads as well,
with sperm produced in the testes of males and
eggs produced in the ovaries of females.
Fertilization in starfish species is
normally done externally with both sperm and
egg being released into the environment unlike
humans who have specialized tracts for
reproduction internally. Starfish gametogenesis
is often influenced by environmental factors
such as temperature and light. In humans,
gametogenesis is regulated by hormonal signals
from the hypothalamus, pituitary gland, and
gonads. This also means that unlike starfish
humans do not have a timing for when to
reproduce and instead rely on deeper emotional
and psychological responses and this may
explain the constant need to produce sperm and
menstruation lasting up until 50 years old for
most females. While both starfish and humans
undergo gametogenesis to produce gametes,
there are significant differences in the process
and characteristics of gametes due to the
different reproductive strategies and
evolutionary paths of these organisms.
The appearance of starfish ovaries can
be quite distinct and unique. They typically have
a granular or lobular appearance, often
resembling a cluster of small beads strung
together. This appearance is primarily due to the
structure and organization of the ovaries within
the starfish reproductive system. In starfish, the
ovaries are part of the gonads, which are
responsible for producing eggs for reproduction.
The ovaries contain numerous follicles, each of
which houses developing eggs at various stages
of maturation. These follicles are arranged in
clusters or lobules throughout the ovarian tissue.
The bead-like appearance of the ovaries arises
from the aggregation of these follicles. Each
follicle contains a single developing egg
surrounded by supportive cells. As the eggs
mature, they grow in size and become more
prominent within the follicles. This growth and
clustering of developing eggs give the ovaries
their characteristic appearance. Additionally, the
ovaries are rich in blood vessels and connective
tissue, which further contribute to their texture
and appearance. These structures provide the
necessary nutrients and support for the
developing eggs as they mature within the
ovaries.
In contrast, human ovaries are typically
almond-shaped and about the size of a walnut.
This shape allows them to be compact yet
contain numerous follicles and structures
necessary for egg development. The surface of
the ovaries is covered by a layer of epithelial
cells called the ovarian surface epithelium. This
layer can appear smooth but may also have
small protrusions or indentations, giving the
surface a slightly bumpy texture. Similar to
starfish ovaries, human ovaries contain follicles
that house developing eggs. These follicles are
embedded within the ovarian tissue and vary in
size and appearance depending on the stage of
development of the contained egg. Immature
follicles are smaller and more numerous, while
mature follicles are larger and more prominent.
The ovaries consist of an outer cortex and an
inner medulla. The cortex contains the follicles
and is where oocyte (egg) development occurs.
The medulla contains blood vessels, lymphatic
vessels, and nerves that supply the ovary with
nutrients and support its function. Overall, the
appearance of human ovaries reflects their
intricate structure and role in the human female
reproductive system.
In starfish, the male reproductive
organs, called testes, have a distinctive
appearance that reflects their function and
evolutionary adaptations. Starfish testes are
typically whitish in color and can be found in the
arms or rays of the starfish, often located near
the central disc. The appearance of starfish testes
is largely due to their internal structure and the
Developmental Zoology
arrangement of reproductive cells within them.
Unlike mammals, starfish do not have distinct,
encapsulated testes. Instead, their testes consist
of numerous branching tubules or follicles that
are interwoven with connective tissue and
muscle fibers. Within these tubules, specialized
cells called spermatogonia undergo a series of
developmental stages to produce sperm cells. As
sperm cells mature, they accumulate within the
tubules, eventually being released into the
surrounding seawater during spawning events.
The branching tubular arrangement of starfish
testes maximizes the surface area available for
sperm production, enabling these animals to
produce large quantities of sperm to increase the
likelihood of successful fertilization during
spawning events.
Human testes have a complex internal
structure, consisting of seminiferous tubules
where sperm production occurs, surrounded by
supporting tissue and vasculature. They have a
relatively uniform oval shape. The testes often
have smooth appearance due to the clustering of
cells. The testes are composed of various tissues,
including seminiferous tubules, interstitial
tissue, blood vessels, and connective tissue.
Human sperm are distributed throughout the
different tubes found in the testes and often form
clusters that vary in the recency of their
production spreading outward to more mature
sperm. In both humans and starfish,
spermatogenesis is supported by various types of
somatic cells. In humans, Sertoli cells within the
seminiferous tubules provide physical and
nutritional support to developing sperm cells.
Leydig cells in the interstitial tissue produce
testosterone, which is essential for
spermatogenesis.
In contrast to Amniotes (mammals,
birds, and reptiles) where spermatogenesis
occurs in seminiferous tubules. Anamniotes
(amphibians and fish) spermatogenesis takes
place in a structure formed when Sertoli cells
engulf a spermatogonia stem cell, called cysts.
However, spermatogenesis in amphibians is
divided also into prespermatogenesis and active
spermatogenesis also known as spermiogenesis,
similar to humans.
Prespermatogenesis covers the earliest
stages of germ cell proliferation and their
entrance into the first meiotic cycle in
developing testes. This early phase starts during
fetal life, right after primordial germ cells
(PGCs) invade the genital ridges of the forming
gonads and become gonocytes, also known as
prespermatogonia or prospermatogonia. These
gonocytes start mitotic divisions (M gonocytes),
giving rise to a pool of transient (T) gonocytes
which are dormant until they become stem cells
(SSCs) during puberty. SSCs are mitotically
active and can either renew the adult
spermatogonial pool or give rise to several
generations of secondary spermatogonia which
will enter meiosis, resulting in sperm production
or spermatogenesis (Haczkiewic et al., 2017).
During spermatogenesis in anurans, the
gonocytes proliferate in the developing testes of
tadpoles which are surrounded by pre-Sertoli
cells.
One of the notable characteristics of
cystic spermatogenesis is the transient germinal
epithelium which requires the turnover of Sertoli
cells in adults. This is also the reason as to why
anamniotes have higher spermatogenic
efficiency and Sertoli-supporting capacity
compared to amniotes. These somatic cells can
divide when in contact with mitotically active
spermatogonia stem cells forming new cysts
periodically. They differentiate and proliferate as
the cyst grows and matures. The number of
Sertoli cells that form a given cyst grows along
with the mitotic division of the germ cells. These
cells are responsible for providing the right
environment for germ cell proliferation, meiosis,
and sperm differentiation.
Developmental Zoology
The spermatogonial stem cells are
enclosed within cysts formed by Sertoli cells.
All the germ cells contained in one cyst are at
the same developmental stage as a consequence
of incomplete cytokinesis. Cytoplasmic bridges
keep these germ cells connected. SSCs can
either proliferate (self-renew) or produce
secondary spermatogonia that will enter meiosis
(primary and secondary spermatocytes),
producing round spermatids and then sperm
(Roco et al., 2021). This signifies the second
phase known as spermiogenesis or active
spermatogenesis. Once the spermiogenesis has
finished, the cysts open to release spermatozoa
in a process called spermiation. The basal parts
of the Sertoli cells in anurans remain after, then
later regenerate the Sertoli cells.
Similarly, oogenesis in amphibians also
has unique mechanisms that differ from other
vertebrates. Early germ cell differentiation into
primary oocytes in most mammals occurs in
fetal life, whereas, amphibian germ cells enter
differentiation into oocytes continuously,
throughout their life. The amphibian oocyte
increases in size during folliculogenesis due to
the incorporation of the yolk protein
vitellogenin. Only a portion of the developing
oocytes reach final maturity and the remaining
parts undergo atresia. To control fecundity, the
balance between the numbers of mature and
atretic oocytes is important in frogs and fish
(Safholm et al., 2012).
Moreover, Hemiphractidae (marsupial
frogs) exhibit multinucleated oogenesis, a
modification far from the common vertebrate
characteristic of a single nucleus per oocyte
(mononucleate oocyte). It is considered to be a
derived character and this shift from
mononucleated to multinucleated oogenesis has
occurred several times during their evolution
(Pino, 2018). In small oocytes, nuclear size is
uniform but as the oocyte grows, the germinal
vesicles (oocyte nuclei) located toward the
periphery are enlarged. On the other hand, in
larger oocytes, peripheral nuclei enlarge and
chromosomes acquire the lampbrush
configuration which is a distinct characteristic of
amphibian oocytes. In contrast, the nuclei
located in the central part of the oocyte remain
small and eventually disappear. Most nuclei
become small and gradually disappear at the
beginning of vitellogenesis. However, the
remaining germinal vesicles enlarge further
before degenerating. Eventually, only one
nucleus remains as the germinal vesicle of the
oocyte. There are no morphological
characteristics that identify the nucleus that will
remain as the germinal vesicle during the
multinucleate period (Pino, 1989).
The multinucleate condition can be
derived from various mechanisms. It can be
from mitotic divisions of oogonia within an
ovarian cyst, followed by cell fusion as in
Flectonotus pygmaeus. The multinucleated
oocyte can also be a result of three oogonial
divisions without cytokinesis just like in
8-nucleated Ascaphus truei.
In some species, the female routinely
produces hundreds or thousands of eggs at a
time, such as sea urchins and frogs. These eggs
are derived from oogonial stem cell populations
which are self-renewing and can generate a new
cohort of oocytes routinely throughout the
organism’s lifetime. On the contrary, in other
species such as humans and most mammals,
only a few eggs are produced during the lifetime
of an individual because the oogonia divide to
form a limited number of egg precursor cells
(Gilbert, 2000).
Another contrast to the gametogenesis
of humans , amphibians and echinoderms is the
gametogenesis of avians. Oogenesis in avians is
also composed of vital stages to produce a viable
haploid egg cell capable of becoming an
embryo. The development of this embryo, like
Developmental Zoology
some amphibians, occurs outside the female’s
body (Wake & Dickie, 1998).
Oogenesis also begins with the
Primordial Germ Cells (PGCs) which will
undergo specialization either through epigenesis
as observed in humans and salamanders or via
hereditary mode occurring in birds, frogs, fish
and insects. During PCGs migration to the
gonads, it will undergo epigenetic
reprogramming, which includes dynamic
changes in histone modifications and
genome-wide DNA demethylation (Choi et al.,
2022; Rengaraj et al., 2011). PGCs polarize
initially during ovo embryonic development, and
they passively migrate toward the anterior
region as a result of the embryo's morphogenetic
movement. PGCs enter blood arteries through
their lumina and stay in the bloodstream entering
the genital ridge at approximately at 2-day old
embryo (Kang et al., 2015).PGCs proliferate and
differentiate into oogonia which undergoes
meiotic division and halt on prophase I.
Luteinizing hormone (LH), along with the
accumulation of the yolk promotes follicle
maturation up until the rupture of follicle wall,
releasing the ovum (Lohmann breeders, 2020).
In chickens and other bird species,
occurs asymmetric gonadal development—in
which the female's left gonad only develops into
an effective ovary while the right gonad
degenerates (Yu et al., 2017) contrary to other
animal classes which have multiple functional
ovaries. Unlike amphibians which have seasonal
reproduction depending on phenological
variation (Silla et al., 2021), hens have
numerous primordial cells, one follicle matures
per day and is released to the oviduct regardless
of fertilization. Follicle stimulating hormone
(FSH) receptor expression is thought to be
inhibited by inhibitory signals, which are
released from the pituitary gland. Individual
follicles that lose this inhibitory signal express
FSH receptors, triggering the synthesis of steroid
hormones and other factors necessary for the
survival of follicle-associated cells (Lohmann
breeders, 2020).
Meanwhile a complex regulatory
network drives spermatogenesis, and recent
research has revealed that gene changes and
signaling pathways can impact spermatogenesis
through controlling the production of
spermatogonial stem cells. (SSCs) (Hu et al.,
2022b). SSCs, initiators of spermatogenesis are
not altered during the process. Spermatogonia
undergo meiosis resulting in primary
spermatocytes becoming secondary
spermatocytes, then spermatids and ultimately,
spermatozoa (Hu et al., 2022; Lan et al., 2011;
Tagami et al., 2017).
While this process takes 30-78 days
across various species under the normal
conditions (Hess & França, 2009), birds’
spermatogenesis appears much faster with a
duration of 11-14 days specifically for non
passerine birds (Noirault et al., 2006). This
could be an act of adaptation against sperm
competition where the testes would produce
maximum sperm output but a study of Lüpold et
al. (2011) revealed that one species discards
spermatids with unfavorable characteristics and
let sertoli cells supported only a few number of
round spermatids towards their maturation,
leaving the rest arrested on the said stage. This
screening process is an evolved useful
modification to optimize the quantity and quality
of sperm.
The irregular size of the chicken’s testes
with the left testis is significantly larger and
elongated compared with the right testis
throughout the growth of the chickens (one week
to three months old). Both appear to be smooth
and creamy. However, on the 4-month old
chickens, testes are more bean-shaped than
elongated. This growth in size causes them to
occupy large proximity extending to the lungs
Developmental Zoology
and an increase of the Gonadosomatic Index
(GSI) in the third month in which the chickens
are identified to be sexually mature. Compared
to mammals, testes of chickens are located in the
abdominal cavity where high body temperature
is achieved to support the process of
spermatogenesis (Mfoundou et al., 2022). This
temperature is one of the contributing factors
why chickens have been selective breeders,
contrary to other animals such as mammals
which do not follow the season for breeding
(Cheng, 2010).
In the first week of the development of
the chicks the seminiferous tubules (ST)
exhibited resorption by apoptosis. Some cease
growing but normal ST demonstrates regression
of degenerating cells within the apoptotic
tubules. Many factors regarding migration,
adhesion, proliferation, and survival of PGCs
influence this process. In the first month begins
the clearer formation of the ST,supporting the
differentiation of the spermatogonia to the
primary spermatocytes. This age is crucial to
the development of the gonads. From here,
continuous growth, development together with
vascularization will occur as PGCs of chickens,
unlike those in other animals, are transported
from avian embryos to the future gonadal region
by blood circulation, where they multiply
concurrently with the gonads' expansion
(Nakamura et al., 2013). Spermiogenesis will
then take place, equipping the spermatids for the
penetration of the egg through chemical and
physical modifications specifically acrosome
formation (Mfoundou et al., 2022) which is a
key component lacking from amphibian sperm
(Kuehnel & Kupfer, 2012).
CONCLUSION
The comparative analysis of
gametogenesis across chickens, frogs, humans,
and starfish unveils a captivating mosaic of both
shared and distinct features in hormonal
regulation, timing, and evolutionary adaptations.
Testosterone, estrogen, and follicle-stimulating
hormones emerge as pivotal regulators,
exhibiting variations in their functions and
pathways across species. Furthermore, the
timing of gametogenesis displays diverse
patterns, ranging from continuous activity in
chickens to seasonal cues in frogs, cyclic
processes in humans, and environmental
influences in starfish. Amidst these variations,
however, lies a remarkable degree of
conservation in the fundamental principles of
germ cell development, meiosis, and gamete
formation. This conservation underscores the
deep evolutionary roots of sexual reproduction,
emphasizing the essential role of gametogenesis
in perpetuating species diversity. Through
comparative studies, we unravel the intricate
interplay between adaptation and conservation,
shaped by evolutionary pressures and
environmental factors. Such insights not only
enrich our understanding of reproductive
biology but also inform conservation strategies
and medical interventions, bridging the gap
between fundamental research and practical
applications in addressing reproductive health
challenges across species boundaries.
 Developmental Zoology

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