Concepts of Animal Embryology

Alysha Marie Echano

One of the most important mechanisms in every living organism is evolution,

which led to the differentiation and survival of many species on Earth. Evolutionary
embryology deals with how embryos of different animals were once similar to each
other, but as development occurs, defining features start to appear and depart from
the similarities. The highest degree of resemblance was showed in the phylotypic
stage of embryos. Interestingly, von Baer’s understanding of embryology provided a
better overview of how embryos develop. Thus, it reveals the differences and
similarities of species driven by the process of evolution. Furthermore, the laws of
embryology together with the concept of the phylotypic stage form a framework to
connect the ideas of how changes in the mechanisms of embryonic development
trigger evolution.

According to von Baer, there are four laws governing embryology, and twentieth-
century developmental biologists supported these laws. Oppenheimer and Gould
noted that von Baer described animal development as epigenetic, meaning that an
organism's behavior and environment affect the way its genes work. Also, it is
described as progressing in a branching manner(Barnes, 2014). The first law states
that the general characters within an animal group occur earlier than special ones.
This means that the general structures of a similar animal group tend to appear in the
early stages of development, and later, as ontogeny proceeds, special structures
arise.

The second law states that the less general forms arise from the most general forms
until most distinct forms develop. It justifies that an organism begins as a uniform and
non-complex structure, and develops into a complex and different form. Additionally,
the third law states that an embryo will never pass through the adult form of other
animal rather diverge from them. It explains that every animal embryo looks similar to
each other even if they are distantly related; however, becomes completely dissimilar
in their adult forms. Finally, the fourth law states that an embryo of higher animal
resembles only other embryo but never the other animal. It suggests that the embryo
of a complex organism will never look like the adult form of a less complex organism.

Embryology is an evidence of evolution, and von Baer’s laws are essential in

constructing evolutionary patterns. The four laws help describe evolution by
observing the similarities of embryos and how each became different. Homologous
structures seen in embryos represent the general forms. Later, these structures
disappear and are morphologically modified into different structures with different
functions or no function at all as the embryo develops.

For example, the presence of tail in embryos gradually reduces into a tailbone(a
vestigial structure) in humans and becomes tails(have numerous functions) in other
species. With this, homology supports evolution and suggests that species share a
common ancestry. Hence, the four laws which focus on ontogeny were able to create
a link toward phylogeny, which is the history of the evolution of species, and the
relationship among organisms in line with their common descent. Essentially, the
laws postulated by von Baer are key to the devising of his idea of a progressive
divergence pattern that looks like a funnel.

Moreover, the phylotypic stage plays a crucial role in development and evolution.
This concept is defined as the time point in animal development where species
closely resemble each other. It is also interpreted as the organogenetic period, where
primordial undifferentiated organs are appearing in vertebrates(Richardson, 2012).
Furthermore, it represents the time when highly conserved genes involved in pattern
formation dominate the transcriptome as according by Duboule(Richardson, 2012).
According to a research article, the phylotypic stage has major characteristics
proposed by morphological studies, namely the pharyngula stage, early somite
segmentation stage, and the tailbud stage(Irie & Sehara-Fujisawa, 2007). Ultimately,
this concept is greatly useful in explaining the conserved pattern of limb development
in tetrapods, general developmental mechanisms, and how natural selection acts on
the developmental stages(Richardson et al., 2003).

Importantly, the phylotypic stage is one of the famous derivations from von Baer’s
laws wherein the third law was used as a basis for devising this concept since it
reflects the idea of the divergence of species. The phylotypic stage serves as a
platform where diversification starts and is conserved by the developmental
constraints ensuring the stabilizing selection(Schmidt & Starck, 2004). This concept
is widely accepted in the field of evolutionary developmental biology because of its
massive contribution to the formulation of the phylotypic stage that provides the
source for the body plan of animals within a phylum. Recent transcriptome-based
studies argued that the most conserved stage of development occurs in the mid-
embryonic stage(Uesaka et al., 2022). Thus, this new idea disproved the funnel
development model or known to be the early conservation model, which believed that
the high conservation stage is at the earliest stage of embryogenesis(Irie & Kuratani,
2011).

In vertebrates, the phylotypic stage is still to determine, and in the process of

identifying whether there is an embryonic stage that represents their phylotypic stage.
However, in a study by Irie and Kuratani in 2011 showed that the potential vertebrate
phylotypic period is the phyrangular stage. The study utilized gene expression
profiling through their transcriptomes to identify the conserved stages of the four
vertebrate species namely the mouse, chicken, X. laevis, and zebrafish. All other
vertebrates seem to have highly conserved transcriptomal similarity in their
pharyngula stage except for the X. laevis which corresponds more to the tailbud
stage. The pharyngular stage is characterized to have four distinct features which
include the notochord, dorsal hollow nerve cord, post-anal tail, and series of paired
branchial grooves or gill slits(Hazkani-covo, 2005). Consciously, the study pointed
out that although the pharyngula stage is the best candidate to be the phylotypic
stage of vertebrates, this does not entail that all vertebrate embryos undergo identical
embryonic stages in terms of their morphology.

Despite being the long unchallenged laws of von Baer and the increasing influence of
the phylotypic stage, these concepts faced criticisms as new advance morphological
and genome-based studies come to light. The progressive divergence model derived
from von Baer’s law competed with the findings of Siedel, Sander, and Elinson who
emphasized that the earlier stages of development such as patterns of cleavage
have divergent morphological patterns; thus, do not align with the law. Similarly, the
phylotypic stage remains controversial because of the different interpretations of this
concept. Ballard first proposed the pharyngula stage as the phylotypic while Wolpert
proposed that it was the early somite segmetation stage, and Slack proposed the
taibud stage. Also, the existence of this concept was questioned because of the
obstacles in evaluating constrained developmental stages through morphological
resemblance(Irie and Fujisawa, 2007).

One key problem of phylotypic stage, whether to accept it or not is the heterochrony.
It is the variations of the relative rate or timing of developmental events(Irie and
Fujisawa, 2007). It is also described as a change in time or order of developmental
events that can lead to a change in morphology(Abzhanov, 2013). An example of
heterochrony is observed in the pharyngeal arch formation and somitogenesis
wherein these stages appear almost at the same in mouse but somitogenesis occurs
earlier in zebrafish. As a result, it is hard to determine what embryonic stage is the
vertebrate phylotypic stage, and whether each vertebrate has a phylotypic stage.
Additionally, the term ‘phylotypic period” should be used instead of the phylotypic
stage to prevent the idea that it occurs at a single time point(Hazkani-covo, 2005).
This is because, in vertebrates, the phylotypic stage consists of a broad range of
embryonic morphologies(Schimdt, 2004).

Due to the flaws realized in von Baer’s laws and the phylotypic stage, the hourglass
model was proposed to be an expansion and a revised version of the previous
concept. This was proposed by Denis Duboule and Rudolf Raff in the 1990s by
stating that ontogeny starts with diverse morphologies, and is followed by a
constrained stage with reduced embryonic variations then, proceeds like a von
Baer’s progressive divergence(Hazkani-covo, 2005). The phylotypic stage represents
the narrowest portion or the bottleneck of the hourglass model, and it implies that
evolutionary conservation resides in the mid-embryogenesis thus, reflecting an
hourglass. Interestingly, it is also suggested that evolutionary divergence lies in the
early and late stages of development. To further verify the reliability of the phylotypic
stage on the hourglass model, aside from morphological evidences, researchers
include molecular data that found that there is an involvement of genes, specifically
the Hox genes. As a result, they came up with an insight that genes and expression
patterns have something to do with the hourglass pattern of divergence(Hazkani-
covo, 2005).

It is fascinating to know about the development of animals and the concepts behind
evolutionary developmental biology. With von Baer’s laws, developmental biologists
were able to unfold the mechanisms behind evolution and the forces that influenced
evolutionary conservation, which is a driver of the hourglass divergence of organisms.
Connectedly, the phylotypic stage demonstrates how morphological resemblance is
affected by the conserved patterns of gene expression also affected by natural
selection. Unfortunately, even in the modern evo-devo, the two main concepts in this
paper still need to be refurbished and clarified by further in-depth and detailed
studies. As previous developmental studies have stated, it is not clear whether the
phylotypic period can represent the basic body plan at a higher taxonomical level, as
transcriptome data of a wide range of species is limited. Regardless, our
understanding of the evolution and development relation is remarkably astonishing
as this is vital in developing new concepts or theories to trace our evolutionary history
and the degree of organisms’ relatedness.
Reference(s)

Irie, N., & Kuratani, S. (2014). The developmental hourglass model: a predictor of the
basic body plan? Development, 141(24), 4649–4655. doi:10.1242/dev.107318

Hazkani-Covo, E., Wool, D., & Graur, D. (2005). In search of the vertebrate
phylotypic stage: A molecular examination of the developmental hourglass
model and von Baer’s third law. Journal of Experimental Zoology Part B:
Molecular and Developmental Evolution, 304B(2), 150–
158. doi:10.1002/jez.b.21033

Abzhanov, A. (2013). von Baer’s law for the ages: lost and found principles of
developmental evolution. Trends in Genetics, 29(12), 712–
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Uesaka, M., Kuratani, S., & Irie, N. (2022). The developmental hourglass model and
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https://doi.org/10.1002/jez.b.23027

Richardson, M. K. (2012). A Phylotypic Stage for All Animals? Developmental Cell,

22(5), 903–904. doi:10.1016/j.devcel.2012.05.001

Schmidt, K., & Starck, J. M. (2004). Developmental variability during early embryonic
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Irie, N., & Kuratani, S. (2011). Comparative transcriptome analysis reveals vertebrate
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Irie, N., Sehara-Fujisawa, A (2007). The vertebrate phylotypic stage and an early
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