SIM - Bio 322

College of Arts and Sciences Education
General Education – Science

2nd Flr. DPT Building, Matina Campus, Davao City
Phone No.: (082)300-5456/305-0647 Local 134
UNIVERSITY OF MINDANAO
Physically Distanced but Academically Engaged
Self-Instructional Manual (SIM) for Self-Directed Learning (SDL)

Course/Subject: Developmental Biology (Bio 332/ L)
Week 1 – 18
Name of Teacher: ______________________________
THIS SIM/SDL MANUAL IS A DRAFT VERSION ONLY; NOT FOR REPRODUCTION

AND DISTRIBUTION OUTSIDE OF ITS INTENDED USE. THIS IS INTENDED ONLY
FOR THE USE OF THE STUDENTS WHO ARE OFFICIALLY ENROLLED IN THE
COURSE/SUBJECT.
EXPECT REVISIONS OF THE MANUAL
Prepared by Jessa Mae B. Patalita

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Table of Contents
Cover Page 1
Course Outline Policies 7
Week 1 to 3: DEVELOPMENTAL ANATOMY AND GENETICS 11
The Cycle of Life 12

The life cycle of Rana pipiens, the Leopard Frog 13
Anatomic Approaches to Developmental Biology 16
Comparative Embryology 16
Evolutionary Embryology 17
Medical Embryology and Teratology 19
Let’s Check 1 21
Let’s Analyze 1 21
In a Nutshell 1 22
Anatomy of a Gene 25
Differential Gene Expression 26
Active and Repressed Chromatin 26
Exons and Introns 27
Promoters and Enhancers 27
Transcription Factors 28
DNA Methylation and the Control of Transcription 30
Differential RNA Processing 30
Control of Gene Expression at the Level of Translation 31
Posttranslational Regulation of Gene Expression 32
Let’s Check 2 33
In a Nutshell 2 34
Week 4 to 5: CELL-TO-CELL COMMUNICATION IN DEVELOPMENT 36
Cell Adhesion 37
Cell Affinity 37
Cadherin and Cell Adhesion 38
Cell Migration 39
Group of Paracrine Factors 41
Juxtacrine Signaling 42
Let’s Check 3 43
In a Nutshell 3 45
Maintaining the Differentiated State 46
The Extracellular Matrix 47
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The Epithelial-Mesenchymal Transition 48

Let’s Check 4 49
In a Nutshell 4 51
Week 6 to 7: FERTILIZATION AND EARLY DEVELOPMENT IN VERTEBRATES 53
Structure of the Gametes 54

Sperm Structure 54
Egg structure 54
External and Internal Fertilization 56
External Fertilization in Sea urchin 56
Internal Fertilization in Mammals 59
Let’s Check 5 61
In a Nutshell 5 62
Cleavage 64
Gastrulation 66
Early Development in Sea Urchin 66
Early Development in Snails 68
Early Development in Tunicates 69
Early Development in Nematodes 70
Early Development in Drosophila 71
Let’s Check 6 72
In a Nutshell 6 73
Week 8 to 9: EARLY DEVELOPMENT IN VERTEBRATES AND LATE 76

EMBRYONIC DEVELOPMENT
Early Amphibian Development 76

Early Development in Fishes 78
Early Development in Birds 79
Early Development in Mammals 82
Let’s Check 7 84
In a Nutshell 7 86
Emergence of Ectoderm 89
Construction of the Central Nervous System 89
Epidermis 93
The Neural Crest 93
Paraxial and Intermediate Mesoderm 94
Paraxial Mesoderm 95
Intermediate and Lateral Plate Mesoderm 95
Endoderm 97
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Let’s Check 8 98
In a Nutshell 8 99
Week 10 to 11: DEVELOPMENT OF THE TETRAPOD LIMB AND SEX DETERMINATION 101
Formation of Limb Buds 102

Proximal-Distal Axis of the Limb 102
Anterior-Posterior Axis of the Limb 103
Formation of Digits and Joints in the Autopod 103
Let’s Check 9 105
In a Nutshell 9 106
Chromosomal Determination of Sex 108
Mammalian Sex Determination 108
Environmental Sex Determination 111
Temperature-Dependent Sex Determination 111
Location-Dependent Sex Determination 111
Mammalian Gametogenesis 112
Spermatogenesis 113
Oogenesis 114
Let’s Check 10 116
In a Nutshell 10 118
Week 12 to 13: POSTEMBRYONIC DEVELOPMENT 120
Amphibian Metamorphosis 121

Insect Metamorphosis 122
Regeneration 127
Genes Involved in Aging 134
DNA Repair Enzymes 135
Aging and Insulin Signaling Cascade 136
The mTORC1 Pathway 136
Chromatin modification 136
Exceptions to the Aging Rule 136
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Week 14 to 15: DEVELOPMENT IN HEALTH AND DISEASE, 141

ENVIRONMENT AND EVOLUTION
Genetic Errors of Human Development 142

Teratogenesis 143
Cancer as a Disease of Development 145
Developmental Therapies for Cancer 146
Role of Environment in Producing Phenotypes 150
Developmental Symbiosis 152
Preconditions for Evolution 159
Deep Homology 159
Mechanisms of Evolutionary Change 160
Developmental Constraints on Evolution 161
Selected Epigenetic Variation 161
Week 16 to 18: PLANT DEVELOPMENT 166
Plant Life Cycle 167

Gamete Production Angiosperm 169
Pollen 170
Ovary 171
Pollination and Fertilization 172
Embryogenesis 177
Dormancy 179
Germination 179
Vegetative Growth in Plants 181
Meristems 181
Root Development 181
Shoot Development 182
Leaf Development 182
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Vegetative-to-Reproductive Transition in Plants 186

Senescence in Plants 188
Online Code of Conduct 191
Monitoring of OBD and DED 192
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Course Outline: BIO 238/L – Developmental Biology
Course coordinator: Jessa Mae B. Patalita

E-mail: jbalbanida@umindanao.edu.ph
Student Consultation: Thru e-mail or LMS
Mobile Phone: 09561744467
Effectivity Date: January 2020
Mode of Delivery: Blended (Online with face-to-face or virtual
sessions)
Time Frame: 108 hours
Student Workload: Expected Self-Directing Learning
Requisites: Bio 101/L, Bio 102/L/Chem 235/L
Credit: 3 units lecture and 2 units laboratory
Attendance Requirements: A minimum of 95% attendance is required at all
scheduled virtual or face-to-face sessions
Course Outline Policy
Areas of Concern Details

Contact and Non-contact Hours This 3-unit Lecture and 2-unit Laboratory course self-
instructional manual is designed for blended learning
mode of instructional delivery such as online
sessions through LMS and scheduled face-to-face
meetings in-campus/on-site or virtual sessions for
reviews and examinations. The expected number of
hours will be 54 for the lecture and 108 for the
laboratory which included the face-to-face or virtual
sessions for reviews, laboratory activities and
reports, assessment tasks and examinations
schedules.
Assessment Task Submission Submission of assessment tasks shall be on the 3rd,
5th, 7th, 9th, 11th, 13th, 15th and 18th weeks of the Sem.
The assessment paper shall be attached with a cover
page indicating the title of the assessment task (if the
task is a performance), the name of the course
coordinator, date of submission, and the name of the
student. The document should be emailed to the
Course Coordinator. It is also expected that you
already paid your tuition and other fees before the
submission of the assessment task. If the
assessment task is done in real-time through the
features in the Blackboard Learning Management
System, the schedule shall be arranged ahead of
time by the course coordinator.
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Turnitin Submission (if necessary) To ensure honesty and authenticity, all assessment
tasks are required to be submitted through Turnitin
with a maximum similarity index of 30% allowed. This
means that if your paper goes beyond 30%, the
students will either opt to redo her/his paper or
explain in writing addressed to the course coordinator
the reasons for the similarity. In addition, if the paper
has reached more than 30% similarity index, the
student may be called for disciplinary action following
with the University’s OPM on Intellectual and
Academic Honesty. Please note that academic
dishonesty such as cheating and commissioning
other students or people to complete the task for you
have severe punishments (reprimand, warning,
expulsion).
Penalties for Late Assignments/ The score for an assessment item submitted after the
Assessments designated time on the due date, without an
approved extension of time, will be reduced by 5% of
the possible maximum score for that assessment
item for each day or part-day that the assessment
item is late. However, if the late submission of the
assessment paper has a valid reason, a letter of
explanation should be submitted and approved by
the Course Coordinator. If necessary, you will also be
required to present/attach pieces of evidence.
Return of Assignments/ Assessments Assessment tasks will be returned to you two (2)
weeks after the submission. This will be returned by
email or via the Blackboard portal. For group
assessment tasks, the course coordinator will require
some or few of the students for online or virtual
sessions to ask clarificatory questions to validate the
originality of the assessment task submitted and to
ensure that all the group members are involved
Assignment Resubmission You should request in writing addressed to the
course coordinator his/her intention to resubmit an
assessment task. The resubmission is premised on
the student’s failure to comply with the similarity
index and other reasonable grounds such as
academic literacy standards or other reasonable
circumstances e.g., illness, accident, or financial
constraints.
Re-marking of Assessment Papers You should request in writing addressed to the
and Appeal program coordinator your intention to appeal or
contest the score given to an assessment task. The
letter should explicitly explain the reasons/points to
contest the grade. The program coordinator shall
communicate with the students on the approval and
disapproval of the request.
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If disapproved by the course coordinator, you can

elevate your case to the program head or the dean
with the original letter of request. The final decision
will come from the dean of the college.
Grading System All culled from BlackBoard sessions and traditional
contact:
For Lectures: 40% of the total grade
Course discussions/exercises – 40%
1st formative assessment – 10%
2nd formative assessment –10%
3rd formative assessment – 10%
All culled from on-campus/onsite sessions (TBA):
Final exam – 30%
For Laboratory: 60% of the total grade
Course Laboratory Report/exercises – 40%
1st formative assessment – 10%
2nd formative assessment –10%
3rd formative assessment – 10%
All culled from on-campus/onsite sessions (TBA):
Final exam – 30%
Submission of the final grades shall follow the usual
University system and procedures.
Preferred Referencing Style Depends on the discipline; if uncertain or inadequate,
use the general practice of the APA 6th Edition.
Student Communication You are required to create a umindanao e-mail
account which is a requirement to access the
BlackBoard portal. Then, the course coordinator shall
enroll the students to have access to the materials and
resources of the course. All communication formats:
chat, submission of assessment tasks, requests, etc.
shall be through the portal and other university
recognized platforms.
You can also meet the course coordinator in person

through the scheduled face to face sessions to raise
your issues and concerns.
For students who have not created their student e-

mail, please contact the course coordinator or
program head.
You can also contact the course coordinator through

the scheduled face to face sessions or social media
for issues and concerns relevant to the course. For
related concerns, you can contact the CASE Dean,
Science Program Head, Library and Guidance Help
Desks.
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For students who have not created their student

email, please contact the course coordinator or
program head.
Contact Details of the Dean Dr. Khristine Marie D. Concepcion
Email:
khristinemarie_concepcion@umindanao.edu.ph
Phone: (082)300-5456/305-0647 Local 134
Contact Details of the Program Head Edgar B. Retorta
Email: edbalre8@gmail.com
Phone: (082)300-5456/305-0647 Local 149
Students with Special Needs Students with special needs shall communicate with
the course coordinator about the nature of his or her
special needs. Depending on the nature of the need,
the course coordinator, with the approval of the
program coordinator, may provide alternative
assessment tasks or extension of the deadline for
submission of assessment tasks. However, the
alternative assessment tasks should still be in the
service of achieving the desired course learning
outcomes.
Well-being Welfare Support Help ZERDSZEN P. RANISES
Desk Contact Details E-mail: gstcmain@umindanao.edu.ph
Phone: (082) 300-5456
Hotline no: 0950-466-5431Phone: (082)300-
5456/305-0647 Local 149
Library Contact Details Brigida E. Bacani
Email: library@umindanao.edu.ph
Phone:0951-3766681
Course Information
CC’s Voice: Hello and welcome to BIO 238/L-Developmental Biology! Every living
organism begins with a single cell that multiply and differentiate to a fully
functional member of an ecosystem. In this subject course, you are going to
explore the molecular principles and concepts underlying this development
in several model organisms. Furthermore, you will be encouraged to observe
development through personalized experiments that will make you
appreciate the complexity and specificity of these processes.
CO: In Developmental Biology, we will discuss the development of plants and

animals, with a particular emphasis on the molecular genetic basis for
developmental events. The entire span of development will be examined,
from the formation of germ cells and fertilization through embryonic
development and even up through post-embryonic development,
senescence, and death.
Let’s Begin!
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Big Picture
DEVELOPMENTAL ANATOMY AND GENETICS
Week 1 to 3: Unit Learning Outcomes (ULOs):

At the end of discussion, you are expected to:
a. Discuss the general pattern of life cycle and anatomical approaches in the
study of Developmental Biology.
b. Discuss the genetic approach in the study of Developmental Biology.
Big Picture in Focus

ULO-a. Discuss the general pattern of life cycle and anatomical approaches in
the study of Developmental Biology.
Metalanguage
In this section, the essential terms relevant to Developmental Biology and to

demonstrate ULO-a will be operationally defined to establish a common frame of reference
as to how the texts work. You will encounter these terms as we go through the lessons in
Developmental Biology. Please refer to these definitions in case you will encounter difficulty
in understanding some concepts.
1. Developmental Biology is the study of the embryonic and developmental process

in multicellular organisms, including plants and animals.
2. Embryology is the study of animal development, specifically at the phase of an
organism that exist between fertilization and birth.
3. Fertilization is the fusion of mature sex cells, the sperm and the egg.
4. Cleavage involves the division of the zygote cytoplasm into small cells called
blastomeres after fertilization. By the end of cleavage, the blastomeres will form a
sphere called blastula.
5. Gastrulation involves the rearrangement of blastomeres. By the end of the gastrula
stage, the embryo will contain three germ layers that will interact to generate organs.
6. Organogenesis is the process of the formation of tissues and organs.
7. Larva is the young version of an organism that requires to undergo metamorphosis
to be a sexually mature adult.
8. Gametogenesis is the process of the formation of gametes.
9. Precursor cells are partially differentiated stem cells that are already committed to
forming a particular type of cell.
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Key Knowledge
Developmental Biology is the discipline of science that deals with the and
embryonic developmental processes in multicellular organisms. This study aims to answer
several questions: (1) how does a single cell give rise to hundreds of different cells? (2) how
can cells in our body organize themselves in functional structures? (3) how can cells regulate
cell division? (4) how are gametes different from other somatic cells? (5) what are stem cells?
(6) how do changes in development create new body forms and how does this influence
evolution? And (7) how does the environment affect the development of some organisms?
There are three ways to approach the study of embryology, or the study of the embryo:
anatomical, experimental and genetic approach.
The Cycle of Life
In general, all animal collectively undergoes the process of embryogenesis, the

development and formation of embryos between fertilization and hatching. There are six (6)
recognized stages in embryogenesis:
Figure 1. Life Cycle of a Frog (Gilbert and Baressi, 2016)
1. Fertilization, a process that involves the fusion of mature gametes, or sex cells. The
fusion of these gametes completes the number of chromosomes that will further
instruct the formation and development of the new individual.
2. Cleavage occurs after fertilization and involves the rapid division of the zygote, the
cell formed in fertilization, forming smaller cells called blastomeres. Blastula, a
sphere of blastomeres, is formed by the end of cleavage.
3. Gastrulation is the series of cell rearrangements, where the blastomeres changes
their position relative to others. By the end of this stage, the embryo will now contain
three germ layers – the ectoderm, mesoderm and endoderm.
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4. Organogenesis occurs after gastrulation and mainly involves the formation of

tissues and organs. The cells that form the tissues and organs are often from different
germ layers. For instance, the epidermis (outer layer of the skin) originates from the
ectoderm, while the dermis (middle layer of the skin) originates from the mesoderm.
Furthermore, cells that are precursors of different specialized cells undergo
migrations from their place or origin to their final location.
5. Metamorphosis applies for organisms that hatches eggs or hatchlings that are not
sexually mature. For them to be able to be mature sexually and undergo reproduction,
they need to undergo metamorphosis. Young, immature organisms are called larva.
6. Gametogenesis is the process of formation of new gametes to produce the next
generation of organisms. Gametes and their precursor cells are collectively called as
germ cells. Body cells are called somatic cells. Eventually, an organism will undergo
senescence and dies with some supporting the early embryogenesis of their offspring
with their nutrients. With the absence of competition, the life cycle is then renewed.
The table below shows the
The Life Cycle of Rana pipiens, the Leopard Frog
The gametogenesis of male and female leopard frog is mainly affected by season
and temperature. Female frogs often develop their ovaries during spring, where photoperiod
and temperature informs the pituitary gland to produce the hormone estrogen. The estrogen,
in turn, instructs the liver to create and secrete yolk proteins for the enlarging eggs in the
ovary. The yolk will be transported to the bottom portion of the egg, called the vegetal
hemisphere. The vegetal hemisphere will serve as the food for the developing embryo. The
other half is called animal hemisphere. Male frogs, on the other hand, creates sperms during
summer with enough sperms produced by fall and before hibernation for spring’s breeding
season.
Figure 2. Eggs and coupling of Rana pipiens (Gilbert and Baressi, 2016)
Fertilization of leopard frogs occur externally. Male frogs grab the female frogs’ back
and fertilizes the egg as the female releases them. Fertilization completes the formation of
the diploid zygote nucleus. In addition, it also allows the cytoplasmic migration within the
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egg that will determine the different axes (anterior-posterior, dorsal-ventral and right-left) of
the frog. Lastly, fertilization activates cell cleavage and gastrulation.
During cleavage, the newly formed zygote divides into thousands of cells, with the
size remaining the same. The cells in the animal hemisphere divides faster than the vegetal
hemisphere. In the animal hemisphere, blastocoel, a fluid-filled cavity, forms which will allow
movements in the rearrangement of cells in gastrulation. Gastrulation begins in the area 180°
opposite to the point of sperm entry, forming a dimple called blastospore. The blastospore
will mark the dorsal side of the embryo. In addition, the dimple will also expand into a ring,
where cells migrating through the blastopore becoming the mesoderm. The cells that remain
on the outer surface and will cover the remaining cells including the vegetal hemisphere will
form the ectoderm and the endoderm will originate from the large, yolky cells in the vegetal
hemisphere. At the end of gastrulation, the mesoderm will be in between the inner endoderm
and outer ectoderm. The ectoderm has the precursor cells from for the epidermis, brain and
nerves; the mesoderm hast the precursor cells for connective tissue, blood, heart, skeleton,
gonads and kidneys; and the endoderm will have the precursor cells for the gut and the
respiratory system.
Figure 3. Cleavage and Gastrulation in Rana pipiens. (Gilbert and Baressi, 2016)
The notochord and the neural tube play an essential part in organogenesis. By this
stage, the embryo is called a neurula. The notochord originates from mesodermal cells
located in the dorsal portion of the embryo. This rod structure signals the ectodermal cells to
form a tube and become a nervous system, rather than form the epidermis. The neural tube
forms by the elongation, stretching and folding into the embryo. These two neural structures
will initiate and signal the other cells to continue and differentiate into different tissues and
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organs. Somites, or precursors of the frog’s back muscles, spinal vertebrae and dermis, will
form from the mesodermal cells adjacent to the neural tube and notochord. Later, the mouth
and anus will form on the embryo, which will further elongate to from a tadpole. Nerve
connections will be slowly established, as well as the gills. The larva will be ready to hatch
after exhaustively feeding from the yolk supplied by the mother frog.
Figure 4. Organogenesis in Rana pipiens.
Figure 4. Organogenesis in Rana pipiens (Gilbert and Baressi, 2016)
The metamorphosis of a tadpole to an adult frog will be initiated by hormones from

the tadpole’s thyroid gland. The new structures evident in a transitioned tadpole to an adult
frog will be its hindlimbs and forelimbs from tadpole paddles, bony skull from cartilaginous
skull, modified mouth, jaw and fly-catching tongue muscle from tadpole’s horny teeth, shorter
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intestine from lengthy, herbivorous intestine, and lungs from regressing gills. At the end of
the metamorphosis, gametogenesis begins. The germ cells in frogs will undergo meiosis.
In leopard frogs, the eggs can take 3 years to mature while the sperm can mature as soon
after metamorphosis. It is essential that metamorphosis is finished before winter sets in. The
figure below shows the life cycle of leopard frogs, emphasizing on anatomical
representations of every stage.
Anatomic Approaches to Developmental Biology
Developmental Biology has been mostly studied using three anatomical approaches:
(1) Comparative embryology, which explores the anatomical changes in embryo
development across organisms; (2) Evolutionary embryology, which studies the changes
and constraints in development that may have cause evolutionary changes in organisms
relative to their ancestors; and (3) Teratology, which studies birth defects.
Comparative Embryology
The first attempt to compare the development across organisms was done by
Aristotle in the 14th century. He had noted variations in life cycles, oviparity, viviparity and
ovoviviparity in his book The Generation of Animals. He also differentiated two major cell
division patterns by which embryos are formed: holoblastic and meroblastic pattern. By 1651,
William Harvey have concluded that all animals come from eggs – even mammals – (Ex
vovo omnia, which means ‘all from the egg’) in his work On the Generation of Living
Creatures. Harvey also was the first to observe the blastoderm, the yolk-free cytoplasm in
chicken egg which will give rise to the embryo, and the ‘islands’ of blood tissue form before
the heart does. He also proposed that amniotic fluid functions as a shock absorber for the
embryo. By 1652, Marcello Malphigi published the first microscopic account of chick
development, where the neural groove, somites and first circulation of veins and arteries
were identified. Both Aristotle and Harvey supported epigenesis, where organs are of the
embryo are formed de novo, or from scratch, at each generation. In contrast, Malphigi
supported the view of preformation, where the organs are believed to be present in
miniature form within the gametes.
Preformationist hypothesis was established by Albrecht von Haller and Charles

Bonnet. According to this hypothesis, organs are already prefigured, and embryonic
development merely required the growth of existing structures, thus, the formation of new
structures is not needed. Another hypothesis was encapsulation, where the germ cells of
adult organisms already have the prefigured state of the next generation. In contrast to von
Haller and Bonnet, Kaspar Friedrich Wolff, a German embryologist, believed in epigenesis
and demonstrated that embryonic parts develop from tissues that have no counterpart in the
adult organism. By 1767, Wolff detailed the formation of the intestinal tube from the folding
of originally flat tissue of a chick embryo, establishing his firm support for epigenesis. Wolff
also indicated a vis essentialis (‘essential force’) that organizes embryonic development,
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though, it was not specified. This essential force was further explored by German philosopher
Immanuel Kant and his colleague Friedrich Blumenbach and he called it Bildungstrieb or
developmental force observable through experimentation. Blumenbach stated that
development can proceed through a predetermined force or instruction inherent in the
embryo. Currently, with the help of the field of Genetics, it is now established that instruction
for development is already present in a fertilized egg.
By 1820s, Christian Pander established the three germ layers – ectoderm,

endoderm and mesoderm. These three layers are the origin of cells that will differentiate to
tissues and specific organ systems. Triploblastic animals have these three layers while
diploblastic animals, such as poriferans and ctenophores, lack a true mesoderm. Heinrich
Rathke was the first one to identify pharyngeal arches that may later differentiate to gills for
fishes and jaws and ears in mammals. Rathke also established the idea of cellular induction,
where tissues interact prior and during differentiation. Karl Ernst von Baer supported
Pander’s theory of germ layer and established his own generalizations (von Baer’s laws)
primarily stating that all vertebrate embryos begin with basically similar structures. These are
the four von Baer’s law:
1. The general features of a large group of animals appear earlier in development than
do the specialized features of a smaller group.
2. Less general characters develop from the more general, until finally the most
specialized appear.
3. The embryo of a given species, instead of passing through the adult stages of lower
animals, departs more and more from them.
4. Therefore, the early embryo of a higher animal is never like a lower animal, but only
like its early embryo.
By the late 1800s, it was established that cells are continuously moving in an embryo
and that they do not have a constant shape. There are two major cells in the embryo:
epithelial cells and mesenchymal cells. Epithelial cells are tightly connected to one another
in sheets or tubes while mesenchymal cells are unconnected and independent units.
Morphogenesis highly depends on the two cells specifically on: (1) direction and number
of cell divisions, (2) changes in cell shapes, (3) cell movement, (4) cell growth, (5) cell
death and (6) the changes in the composition of the cell membrane or secreted
products. In order to track such dynamic movements and changes, tracing of cell lineages
through fate maps were established. Fate maps show the fate of the cell or a group of cells
in the embryo, portraying which parts of the embryo will develop into which tissues. Other
methods that are practiced in tracking cell lineages are through direct observation of living
embryos, dye markings, and genetic labeling.
Evolutionary Embryology
Evolutionary change is often based on developmental change. Without changes in

development, there would be no variations needed to establish evolution. Charles Darwin’s
establishment of the theory of evolution gave new focus to comparative embryology. Darwin
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believed that embryonic resemblance is a strong support for genetic connectedness. He

stated in his work, On the Origin of Species in 1859 that “Community of embryonic
structures reveals community of descent”.
Before Darwin’s revolutionary work, some taxonomists classify organisms according

to similarities in larval form. For instance, J.V. Thompson grouped larval barnacles to
arthropods because of its resemblance to larval shrimp in 1830s. Another example was the
work of Alexander Kowalevski, in which he provided glimpse of a bridge of the relationship
between invertebrate and vertebrates by observing notochord, pharyngeal pouches and
germ layers in tunicates, an invertebrate, that are similar to fishes and chicks in 1871.
Furthermore, Darwin also noted some embryonic structures irrelevant to the adult form of
organisms but show their relatedness to other animal such as eyes in embryonic moles,
pelvic bone rudiments in snakes and teeth in baleen whale embryos. In addition to this,
Darwin also noted that adaptations that contributes to survival and fitness of organisms
develop late in embryos and that the differences between genera become more evident as
development persists.
Source: http://group4cladistics.weebly.com/uploads/5/3/1/7/53179435/119029908.jpeg
Figure 5. Analogous and homologous structures
Homology and analogy are two essential concepts in evolutionary embryology.

Homologous structures are those organs whose underlying similarity roots from being
derived from a common ancestral structure. For instance, the wing of a bird and the forelimb
of human are considered as homologous because both have evolved from the forelimb bones
of a common ancestors. Analogous structures, on the other hand, are organs that have
similar functions across organisms rather than their roots of a common ancestor. An example
of these analogous structures are the wings of a butterfly and of a bat, where both function
for flying, but not from the same common ancestor.
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Medical Embryology and Teratology
Medical embryology and teratology often concern humans and the defects that occur
during developmental stages. Abnormalities may include physical deformities, missing parts
and mutant genes and chromosomes that maybe caused by genetic inheritance or
environmental influence. Abnormalities caused by genetics events are called malformations
and malformations can appear as syndromes, where several abnormalities occur
concurrently. Most of these syndromes are observed and studied in animals with the same
condition. An example of a syndrome is piebaldism caused by the dominant mutation in the
KIT gene that encodes for a protein expressed in neural crest cells and in the precursors of
blood cells and germ cells. A patient affected with piebald syndrome exhibit anemia, sterility,
unpigmented regions in the skin and hair, deafness and the absence of nerves that initiate
peristalsis in the gut.
Source: http://images.shoutwiki.com/gn434/b/b7/Mouse.jpg
Figure 6. Piebaldism in human and mouse
Abnormalities caused by exogenous agents such as chemicals, radiation, viruses or

hyperthermia are called disruptions and these agents are called teratogens. Teratology is
the study of disruptions and the environmental agents that disrupt the normal development.
An example of disruptions is phocomelia, a congenital condition of newborn babies common
to pregnant women who has taken thalidomide which is a mild sedative. Newborn babies
that have phocomelia exhibit deficient or absent long bones of the limbs. In 1965, it was
documented that that thalidomide is teratogenic during 34-50 days after last menstruation.
Thalidomide can cause absence or deficiency of ear components from days to 34 to 38. In
addition, it was observed that malformations in the upper limbs occur before those of the
lower limbs since arms form before legs. With the information of the genes and the teratogens
that affect certain aspects of development, anatomical approach of Developmental Biology
has gained new knowledge overtime.
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Source: https://d3i71xaburhd42.cloudfront.net/94a8cf7d92fb7c94c0effe6a7795a523ab53f653/2-Figure1-1.png
Figure 7. Phocolemia in a human baby
References
Gilbert, S. (2010). Developmental Biology. Ninth Edition. USA: Sinauer Associates Inc.
Urry, L. A., Cain, M. L. 1., Wasserman, S. A., Minorsky, P. V., Reece, J. B., & Campbell, N.
A. (2017). Essential biology. Eleventh edition. New York, NY: Pearson Education, Inc.
Self Help.
You can also refer to sources below to help you further understand the lesson.
Streelman, J. T., & Streelman, J. (Eds.). (2014). Advances in evolutionary developmental

biology. ProQuest Ebook Central https://ebookcentral.proquest.com
Minelli, A. (2003). The development of animal form: Ontogeny, morphology, and evolution.
ProQuest Ebook Central https://ebookcentral.proquest.com
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Let’s Check!
Activity 1. Now, that you know the general pattern for the circle of life and the three fields that
comprise the anatomic approach to Developmental Biology, let us try to check your
understanding of these topics. Match the events and the stages of development. Write your
answers in the space provided:
_____ 1. Formation of specialized tissues and organs a. Fertilization
_____ 2. Distinction of the three germ layers b. Cleavage
_____ 3. Rapid division of cells without affecting cell size c. Gastrulation
_____ 4. Formation of blastula d. Organogenesis
_____ 5. Fusion of gametes e. Metamorphosis
_____ 6. Completion of the number of chromosomes f. Gametogenesis

_____ 7. Formation of blastomeres
_____ 8. Long migrations of cells to their final location
_____ 9. Transformation of larva to a sexually mature adult
_____ 10. Development of gametes
Let’s Analyze!
Activity 1. Now that you are acquainted about the general pattern of life cycle among
organisms and the fields that constitute the anatomical approaches to Developmental Biology,
it is important that you will be able to relate this in your future profession. I will require you to
answer the following questions:
1. What are fate maps and what is its importance?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Give other examples of malformations and disruptions prominent in the Philippines.

Describe the causes of these conditions.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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3. Is the study of Developmental Biology only essential among vertebrates? How about
invertebrates?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What is the importance of Developmental Biology in the field of Taxonomy and

Systematics?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 1. Based on your learnings about the general pattern of life cycle among organisms
and the fields that constitute the anatomical approaches to Developmental Biology, feel free
to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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Question and Answers’ List
Do you have any questions for clarifications?

Questions/Issues Answers
1.
2.
3.
4.
5.
Keywords Index
Life cycle stages Teratology
Developmental Biology Homology

Comparative Embryology Analogy
Evolutionary Embryology Disruptions/Syndromes
Medical Embryology Rana pipiens
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ULO-b. Discuss the genetic approach in Developmental Biology.
Metalanguage

1. Differential gene expression is the cause as to why cells differentiate differently

resulting to an organism comprised of cells with different functions.
2. Genome refers to all the genetic material in an organism.
3. Gene is the basic unit of heredity that codes for a particular gene, either RNA or
protein.
4. Chromatin is made up of a complex of DNA and protein that contains the eukaryotic
chromosomes. Histone is the protein component of chromatin while nucleosome is
the basic unit of chromatin structure.
5. Exons are the regions in a DNA that code for a protein. Introns are intervening
regions in a DNA that do not code for protein.
6. Transcription is the genetic process where the information from the DNA is copied
to a strand of RNA.
7. Nuclear RNA or pre-messenger DNA are original transcription products.
8. Promoters are the sites that binds RNA polymerase to the DNA to start transcription.
9. Enhancer controls the efficiency and rate of the transcription process from a specific
promoter. It is a DNA sequence.
10. Silencers are sequences that act to stop the transcription process.
Key Knowledge
The fertilized egg already has the entire genome, or the blueprint used to make up
the whole organism. One of the questions that arose in Developmental Biology along with
this fact is how cells differentiate and take up specific functions within the organism
considering that they have the same information within the cells. This has been established
and explained by the concept of differential gene expression. The following are the postulates
of differential gene expression:
1. Every cell nucleus contains the complete genome established in the fertilized cell.
This means that all differentiated cell in an individual have identical DNAs.
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2. The genes that are not expressed in a differentiated cell are neither destroyed nor
mutated. The potential of the genes to be expressed remains.
3. In differentiated cells, only small percentage of the genome is expressed.
Furthermore, the portion of RNA synthesized in cells is specific for that cell type.
There are several levels in which gene expression can be regulated: (1) differential
gene transcription, (2) selective nuclear RNA processing, (3) selective messenger RNA
translation and (4) differential protein modification. These processes enable the cells
within an organism to have different structure and function.
Anatomy of a Gene
A gene is a specific sequence in a DNA or RNA and is considered as the basic

physical and functional unit of heredity. A specific gene occupies a fixed location in a
chromosome, a thread-like structure made up of protein and a single molecule of DNA.
Chromatin is a structure within a chromosome that contains the DNA and the protein. The
protein counterpart that accounts for most of the chromosome’s weight is called histone. A
nucleosome is the basic functional unit of chromatin, where less than two turns of DNA is
wrapped around eight histones, known as octamer histones. There are two molecules of
each histones H2A, H2B, H3 and H4. The two loops of DNA contain an approximate of 140
base pairs of DNAs. There are 14 points of contact between the DNA and the histones. The
histone H1 is bound to the 60 or so base pairs of ‘linker’ DNA between the nucleosomes.
Source: https://4.bp.blogspot.com/-IG29_RPm-fU/WB44AFC6pPI/AAAAAAAAF9c/vP6zKHGvQywbOIYHhHXeKBklWnjGxdnqwCLcB/s1600/octamer.png
Figure 8. Anatomy of a DNA and a Histone
The following are the elements of a eukaryotic gene:
1. Promoter region, which binds RNA polymerase to initiate transcription.

2. Transcription initiation site, where the first RNA nucleotide is transcribed. This
site is often called the cap sequence because it represents the 5´ end of the RNA,
which will receive a "cap" of modified nucleotides soon after it is transcribed.
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3. Translation initiation site, which comprise of the codon ATG, which will become
AUG in the mRNA. AUG is the codon that starts translation.
4. The 5’ untranslated regions are intervening sequences of base pairs that can
determine the rate at which translation is initiated.
5. Exon, a sequence that code for a protein.
6. Intron, a set of base pairs that has no coding sequences for proteins.
7. Translation termination codon (ex. TAA) that will become the nonsense or stop
codons in the mRNA.
8. The 3’ untranslated region that has the polyA tail that will be transferred to the
mRNA. The polyA tail confers the stability on the mRNA, allows the mRNA to exit
the nucleus and permits the mRNA to be translated into a protein.
Figure 8. Anatomy of a Gene (Gilbert and Baressi, 2016)
Differential Gene Transcription
Active and Repressed Chromatin

The expression and repression of a particular gene is mostly attributed to the
modification of histones. The nucleosomes are stabilized into tight solenoids by histones,
specifically H1. Due to the conformation of nucleosome by H1, the adjacent nucleosomes
are packed tightly together inhibiting transcription factors and RNA polymerases from gaining
access to the genes. In this situation, the chromatin is said to be repressed.
Source: https://www.researchgate.net/profile/Andreas-Vilcinskas/publication/273578679/figure/fig4/AS:268888092966919@1441119325292/Chromatin-modifications-mediated-by-DNA-methylation-and-histone-acetylation-Gene.png
Figure 10. Histone Acetylation and Methylation
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In molecular terms, the repression and activation of gene expression in histone is

made possible by modifying the tails of histones H3 and H4 with two small organic groups:
methyl (CH3) and acetyl (COCH3) residues. Through histone acetylation or the addition of
negatively charged acetyl groups to histones, by histone acetyltransferases, the basic
charge of lysine neutralizes and loosens the histone, thus, activating transcription. In contrast,
histone deacetylases, stabilize the nucleosomes and prevent transcription. Histone
methylation, or the addition of methyl groups to histones, either activate or repress
transcription. This is made possible by histone methyltransferases and usually depends
on the amino acid being methylated and presence of other methyl or acetyl groups.
Exons and Introns

The presence of exons and introns differentiate eukaryotic and prokaryotic genes.
Exons are regions of DNA that codes for proteins while introns are intervening sequences
that do not. During the transcription process, both exons and introns are copied by the RNA.
The resulting RNA or the original transcription product is called nuclear RNA (nRNA) or
heterogenous nuclear RNA (hnRNA), or pre-messenger RNA (pre-mRNA). Introns are
removed as the nRNA leaves the nucleus, and the remaining exons will be spliced and
covalently bonded with each other so that they will form a single transcript for protein
production.
Figure 11. Splicing of introns to form mature transcript (mRNA) (Gilbert and Baressi,
2016)
Promoters and Enhancers

Promoters and enhancers are regulatory sequences that control where and when a
particular gene will be transcribed. Promoters are the sites where RNA polymerase binds to
the DNA to initiate transcription and they are typically located immediately upstream from the
site where the RNA polymerase initiates transcription. These promoters have the sequence
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called TATA box and has 30 base pairs upstream from the site where the first base is
transcribed. Transcription is only possible with the presence of other proteins: (1) TATA-
binding protein (TBP) and TFIIB that properly place the polymerase on the promoter, (2)
TFIIA and TFIIH that stabilizes the complex, and auxiliary transcription-associated factors
(TAF) that also stabilizes the RNA polymerase on the promoter and enable transcription.
Enhancers, on the other hand, controls where and when a promoter can be used and how
gene product will be made. They are responsible for binding transcription factors that activate
the gene or stabilizing the transcription inhiation complex as mentioned. Furthermore,
enhancers can only activate promoters on the same chromosomes (cis-linked promoters),
thus they are also called cis-regulatory elements. Enhancers can bind several transcription
factors, and these specific combinations of factors allows a gene to be active.
Transcription factors
Transcription factors are proteins that controls the rate of transcription of genetic
information from DNA to messenger RNA, by binding to a DNA sequence. There are several
families of transcription factors that are grouped according to their structure. The table below
shows the major transcription factor families and subfamilies.
Some Major Transcription Factor Families and Subfamilies
Family Representative Some Function

Transcription Factors
Homeodomain:
Hox Hoxa1, Hoxb2, etc. Axis formation
POU Pit1, UNC-86, Oct-2 Pituitary development; neural fate
LIM Lim1, Forkhead Head development
Pax Pax1, 2, 3, 6 etc. Neural Specification; eye development
Basic helix-loop- MyoD, MITF, daughterless Muscle and nerve specification;
helix (bHLH) Drosophila sex determination;
pigmentation
Basic leucine C/EBP, AP1 Liver differentiation; fat cell
zipper (bZip) specification
Zinc finger:
Standard WT1, Kruppel, Engrailed Kidney, gonad, and macrophage
development; Drosophila
Nuclear hormone Glucocorticoid receptor, segmentation
receptors estrogen receptor, Secondary sex determination;
testosterone receptor, craniofacial development; limb
retinoic acid receptors development
Sry-Sox Sry, SoxD, Sox2 Bend DNA; mammalian primary sex
determination ectoderm differentiation
Transcription factors have several functions. As mentioned, the combination of

different transcription bound by enhancers can either activate or repress gene expression.
For example, the association of Pax6, Sox6 and MAF transcription factors in lens cells
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can dissociate nucleosome in that area by initiating histone acetyltransferase. In addition,

transcription factors can also stabilize the transcription preinitiation complex that enable RNA
to bind to the promoter. MyoD, a transcription factor essential for muscle cell development,
functions to stabilize TFIIB which also stabilizes RNA polymerase at the promoter site. MyoD
can also initiate nucleosome remodeling and dissociation through binding with histone
acetyltransferase. Transcription factors also combine to initiate coordinated gene
expression, where cell-specific genes are activated simultaneously. The combination of
Pax6 transcription factor to other transcription factors bound to one enhancer in the lens
activates the gene.
There are three domains in transcription factors: (1) DNA-binding domain, which
recognizes the specific DNA sequence in the enhancer, (2) trans-activating domain, which
activates or suppresses gene transcription, and (3) protein-protein interaction domain,
which allows the activity of the transcription factor to be regulated by TAFs or other
transcription factors. In, MITF, a transcription factor essential for ear development and
pigment, the DNA-binding domain is close to the amino-terminal end of the protein and
contains numerous basic amino acid that make contact with the DNA. The trans-activating
domain of MITF has a long stretch of amino acids in the center of the protein that binds to a
TAF, p300/CBP which is a histone acetyltransferase enzyme, causing acetylation and
destabilization of the nucleosomes, thus, allowing the genes to be expressed. The protein-
protein interaction domain of MITF allows it to dimerize with another MITF protein, forming a
homodimer that will be able to bind to an enhancer, that in turn, allows it to activate gene
transcription.
The proteins Trithorax and Polycomb families help in retaining the memory of the
transcriptional state during mitosis. Trithorax keep genes active of nucleosomes of active
genes, whereas Polycomb proteins keep the genes in inactivated state, when the
nucleosomes are condensed. Other transcription factors, called ‘pioneer’ transcription factors
are able to penetrate repressed chromatin and bind to enhancers to allow other transcription
factors to gain access to promoters. Examples of these transcription factors are the FoxA1,
Pax7, and Pbx. FoxA1 binds to enhancers that open up the chromatin to allow transcription
factors access to the promoter. Pax7, on the other hand, activates muscle-specific gene
transcription in a population of muscle stem cells by binding to its DNA recognition sequence
and being stabilized by demethylated H3K4 on the nucleosomes. This action initiates histone
methyltransferase that activates transcription. Pbx proteins allow MyoD to bind to DNA
elements to adjacent DNA sequences that recognize the MyoD.
DNA has regulatory elements that actively suppress gene transcription. Silencers,
such as neural restrictive silencer elements (NRSE), prevent the expression of genes by
preventing a promoter’s activation in any tissue except neurons. The protein that bind to
NRSE is a zinc transcription factor is called neural restrictive silence factor (NRSF), that
appears to be expressed in every cells that is not a mature neuron.
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DNA Methylation and the Control of Transcription
DNA methylation and control of transcription allows a specialized cell to remain as it

is even with mitosis. This is made possible by the methylation of promoters of inactive genes
at certain cytosine residues in a DNA. This process stabilizes nucleosomes and prevents
transcription factors from binding. There are two processes as to which DNA methylation can
repress gene expression: (1) by blocking the binding of transcription factors to enhancers by
methylating one cytosine of a DNA, and (2) by recruiting proteins that facilitates methylation
or deacetylation of histones, stabilizing the nucleosomes. MeCP2 is a protein that can bind
to methylated DNA, binding to histone deacetylases and histone methyltransferases,
resulting nucleosomes to form tight complexes with the DNA, thus, preventing transcription
factors and RNA polymerase to bind and initiate gene expression.
During mitosis, DNA methylation and its pattern is passed through the parent cell to
daughter cells, allowing a specialized cell to remain as it is. Aside from histone deacetylases
and histone methyltransferases, MeCP2 also recruits another enzyme called DNA-
methyltransferase-3 (Dnmt3). This enzyme methylates a large region in a sequence,
repressing it. The methylation pattern is then transferred to another enzyme, DNA
methyltransferase-1 (Dnmt1), that recognizes methyl cytosines on one DNA strand and
places methyl groups on the newly synthesized strand opposite to it. This process creates a
pattern of DNA methylation to be maintained during cell division, ensuring that it is stably
inherited by all the progeny of that cell.
Differential RNA Processing
Gene expression can also be regulated during the synthesis of RNA in the process
of transcription and translation. In order for proteins to be synthesized from nRNA, it must
undergo several stages: (1) it should be processed into a mRNA by the removal of
introns, (2) translocated to the cytoplasm from the nucleus, (3) translated by the
protein-synthesizing apparatus and (4) postranslationally modified to be activated.
There are two major ways of regulation of gene expression through differential RNA
processing: (1) by selecting which nuclear transcripts, or instructions, are processed into
cytoplasmic messages and (2) splicing of mRNA precursors into messages to specify
different proteins by using different combinations of potential exons. Nuclear RNA selection
could also control the development of early development in some organisms. For instance,
sea urchin has the same gene sequences of nRNA in blastula, gastrula and adult tissues but
when it comes to the gene sequences of mRNA, some sequences in sea urchin blastula
messenger RNA are absent in gastrula and adult tissue mRNA.
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Control of Gene Expression at the Level of Translation
The control of gene expression at the level of translation can be regulated by the (1)
shelf-life of the mRNA, (2) the selective inhibition of mRNA translation, (3) microRNAs
and (4) the control of RNA expression by cytoplasmic localization. Some mRNA persists
longer than others, and those that do, the more proteins are translated from it. The stability
of the information or message depends on the length of the mRNA polyA tail. The polyA, in
turn, depends on the 3’ untranslated regions (UTR) on the gene sequence. When
experimentally manipulated, some mRNAs have altered half-lives where long-lived
messages decay rapidly and short-lived mRNAs remain longer.
Regulation of gene expression during translation are also known in oocytes. In

oocytes, mRNAs are made and stored in large amounts and will be used only when
fertilization occurs. Some of these mRNAs code for proteins needed for cleavage and those
needed to determine the fate of cells. Ion signals activate these dormant messages. In
addition, the 5’ cap and the 3’ untranslated region also regulate the accessibility of mRNA to
ribosomes. If the 5’ cap and the 3’ untranslated UTR lacks a polyA tail, the message will not
be translated. mRNAs during this stage form circles, with their 3’ end being brought to their
5’ end.
MicroRNAs are small RNAs complimentary to a portion of a particular mRNA, acting

as an antisense RNA. An example of a microRNA was first observed in C. elegans. The lin-
14 mRNA, coded from the lin-4 gene, inhibit the encoding of a transcription factor, LIN-14,
which is important during the first larval phase of C. elegans development especially at later
stages.
Cytoplasmic localization involves the control of RNA translation by changing the

place of RNA expression. This was observed in Drosophila. This regulation is accomplished
by 3’ UTRs. There are three major mechanisms for the localization of an mRNA: (1) diffusion
and local anchoring, where the mRNAs that diffuse freely in the cytoplasm diffuse to the
posterior pole of the Drosophila oocyte, where they are trapped by proteins that reside in
these regions and can activate the mRNA for translation; (2) localized protection, where
mRNAs that code for the heat shock protein hsp83 (protects the Drosophila embryo from
thermal extremes) are protected in the posterior pole in the cell; and (3) active transport
along the cytoskeleton, where mRNAs are transferred to other parts of the cell by binding
to ‘motor proteins’ that travel through the cytoskeleton to their final destination.
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Figure 12. Cytoplasmic Localization of mRNAs (Gilbert and Baressi, 2016)
Posttranslational Regulation of Gene Expression
After a protein is successfully synthesized, there is still no guarantee that it will

become active and will function within the cell. Some proteins need to be directed to their
specific intracellular destinations in order to function while some proteins are needed to
assemble with other proteins in order to form a functional unit. Ions are also needed to bind
with some proteins in order for the protein to be activated.
References
Self Help.
Kondoh, H., & Lovell-Badge, R. (Eds.). (2015). Sox2: Biology and role in development and
disease. ProQuest Ebook Central https://ebookcentral.proquest.com
Moody, S. A. (Ed.). (2014). Principles of developmental genetics. ProQuest Ebook
Central https://ebookcentral.proquest.com
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Let’s Check!
Activity 2. Now, that you know the genetic approaches to Developmental Biology, let us try
to check your understanding of these topics. Match the following events to its corresponding
level of regulation of gene expression:
____1. Selection of nuclear transcripts to process for a. Differential gene
cytoplasmic messages transcription
____2. Histone acetylation b. Selective nuclear RNA
____3. Transcription factors Processing
____4. MicroRNAs c. Selective messenger RNA
____5. Histone methylation Translation
____6. Silencers d. Differential protein
____7. Cytoplasmic localization of mRNA Modification
____8. mRNA shell-life
____9. DNA Methylation
____10. mRNA splicing using different combinations
of potential exons
Let’s Analyze
Activity 2. Now that you are acquainted about the genetic approaches to Developmental
Biology, it is important that you will be able to relate this in your future profession. I will require
you to answer the following questions:
1. What is the relationship of gene expression and cancer?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. State some differences between prokaryotic and eukaryotic differential gene

expression.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. Give some examples of human proteins that needed ions to be activated and become
functional in a cell.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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4. Human β-globin locus is composed of five genes located on a short region of

chromosome 11, responsible for the creation of the beta parts of the oxygen transport
protein, Hemoglobin. It contains the beta globin gene, as well as the delta, gamma-
A, gamma-G and epsilon globin genes. Thoroughly discuss how the beta-globin gene
is expressed from Human β-globin locus with emphasis in the different regulation of
gene expression.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 1. Based on your learnings about the genetic approach to Developmental Biology,
feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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1.
2.
3.
4.
5.
Keywords Index
Genes Transcription
Promoters Genome
Enhancers Differential Expression
Transcription Factors RNA
Translation DNA
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Big Picture
CELL-TO-CELL COMMUNICATION IN DEVELOPMENT

a. Discuss cell adhesion, cell migration, cell signaling, paracrine factors and
juxtacrine signaling.
b. Discuss the process of maintaining the differentiated state, and the role of
extracellular matrix and the epithelial-mesenchymal transition.

ULO-a. Discuss cell adhesion, cell migration, cell signaling, paracrine factors and
juxtacrine signaling.
Metalanguage
In this section, the essential terms relevant to cell-to-cell communication and to

1. Morphogenesis is the formation of organized animal bodies.

2. Ligands are signaling proteins that are secreted from a cell and are designed
to communicate in response in another cell.
3. Receptors are the binding sites for the ligands during cell-to-cell
communication.
4. Cell migration refers to the movement of cells especially during early embryonic
development.
5. Cadherin calcium-dependent adhesion molecules are major molecules involved
in cell adhesion.
6. Induction is the kind of interaction of two or more cells or tissues of different
histories and properties at close range.
7. In juxtacrine signaling, the proteins from the inducing cell interact with receptor
proteins of adjacent responding cells without diffusing.
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Key Knowledge
Morphogenesis, or the formation of organized animal bodies, is achieved by the

vital process of cell-to-cell communication. In this section, you are going to discover the
answers for these questions: (1) how are separate tissues formed from a population of cells?
(2) how are organs constructed from tissues? (3) how do organs form in particular locations
and how do migrating cells reach their destinations? (4) how do organs and their cells grow,
and how is their growth coordinated throughout development? And (5) how do organs
achieve polarity?
Cell-to-cell communication is mostly achieved by the informational molecules that are

secreted or positioned in the cell’s plasma membrane. Proteins that are secreted from a cell
and are designed to communicate a response in another cell are called signaling proteins,
or ligands. These informational molecules bind to receptors of neighboring cell to set off a
cascade of intracellular reactions resulting to changes in gene expression, enzymatic
activities, and cytoskeletal arrangements, thus affecting cell fate, cell shape and cell behavior.
A receptor in the membrane of one cell that binds the same type of receptor in another cell
represents a homophilic binding, while heterophilic binding involves different receptor
types. In this section, you will learn more about cell adhesion, cell migration, and cell signaling.
Cell Adhesion
Cell affinity
Experimental morphogenesis by Townes and Holtfreter (1995) established the

concept of cell affinity during early embryonic development. Cell affinity ensures that cells
from different germ layers would not get mixed with each other even though they are
experimentally reaggregated with one another. Their experiment has two major results: (1)
reaggregated cells become spatially segregated, that is, each type of cell sorts itself into its
own region when mixed, and (2) the final positions of the reaggregated cells reflect their final
positions in the embryo. These results were possible because of the selective affinity of
cells. For example, the inner surface of the ectoderm has a positive affinity for mesodermal
cells, and a negative affinity for endodermal cells, while the mesoderm has both positive
affinities to ectodermal and endodermal cells. This can also be observed during the first
stages of organogenesis such as in neural plate cells, axial mesoderm cells, and epidermal
cells. However, selective affinities between cells changes throughout development since
cells need to continuously interact with each other especially during morphogenesis.
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Figure 13. Sorting out and reconstruction of spatial relationships in aggregates of

embryonic amphibian cells (Gilbert & Baressi, 2016)
The force that allows these cells to sort themselves in a pattern were first explained
by Malcom Steinberg (1964) through the differential adhesion hypothesis, which mostly relies
on thermodynamic principles. In his experiment, he showed that certain types of cells migrate
centrally when combined to with some cell types but migrate peripherally when combined in
others. Furthermore, Steinberg concluded that cells will rearrange themselves into the most
thermodynamically stable pattern. For instance, if cell types A and B have different strengths
of adhesion, and if the strength of A-A connections is greater than the strength of A-B or B-
B connections, the A cells will sort and become central. In contrast, if the strength of A-A
connections is less than or equal to the strength of A-B connections, then the aggregate will
remain as a random mix of cells. This scenario suggests that early embryo can be seen as
a mixture of aggregated cells until changes in gene expression modify cellular surface
components that will determine cell adhesion strength.
Cadherin and cell adhesion
The major molecules involved in cell adhesion are called calcium-dependent

adhesion molecules, or simply called as cadherin. Cadherins are vital in establishing and
maintaining intercellular connections. Furthermore, they also play an important role in the
spatial segregation of different cell types and morphogenesis.
Cadherins are anchored within cells through catenins. Cadherins have three major
functions: (1) they adhere cells together, (2) link and help assemble the actin
cytoskeleton, thus, making the tubes for mechanical forces, and (3) signal to change
gene expression. For instance, the cadherin-catenin complex forms the classic adherens
junctions that aid in holding epithelial cells together. In addition, the complex also integrate
the epithelial cells into a mechanical unit by binding to actin cytoskeleton of the cells.
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Example of cadherins is the E-cadherin, P-cadherin, N-cadherin, R-cadherin and

protocadherins. E-cadherin is expressed on early mammalian embryonic cells and is later
restricted to epithelial tissues of embryos and adults. P-cadherin allows the placenta to stick
to the uterus. N-cadherin is expressed in cells of the developing central nervous system and
may play a role in mediating neural signals while R-cadherin is vital in the formation of retina.
Protocadherins lack the capability to form a complex with catenins. Similar protocadherins
keeps migrating epithelial cells together while dissimilar protocadherins separate tissues.
Typically, the strengths of these bonding may be achieved quantitatively (the more cadherins
on the opposing cell surfaces, the tighter the adhesion) and qualitatively (the ability of
cadherin types to bind with each other).
An example of the effect of cadherin expression in initiating certain development

events can be seen in the development of legs in chicks. The N-cadherin in chick embryo
appears in the mesenchymal cells of the leg before these cells condense and form nodules
of cartilage, which are the precursors of the limb skeleton. The N-cadherin will not be present
in these cells prior and after the condensation process, thus, insinuating that cartilage
formation in the chick limb is signaled by the appearance of N-cadherin. Another example of
cadherins working together is the attachment of the fertilized egg with the uterus made
possible by E- and P- cadherins.
Cell Migration
Cell migration refers to the movement of cells especially during gastrulation, where
the three germ layers are formed and the first stage of organogenesis. The first stage of
migration is polarization, where the cell identifies its front and back side. Polarization is
initiated by diffusing signals or signals from the extracellular matrix. The signals reorganize
the cytoskeleton of the cell, making the front and back side of the cell structurally different.
The second stage is the protrusion of the cell’s leading edge, mechanized by the
polymerization of the actin filaments at the cell membrane to create long parallel bundles
(filopodia) or broad sheets (lamellipodia). The Rho G-proteins in the cell membrane will
activate the WASP-N protein, nucleating the actin and connecting it to the cadherins and cell
membrane.
The third stage is the adhesion of the cells to its extracellular substrate, which is
highly influenced by integrin proteins located in cell membrane. Integrin attaches to actin to
form focal adhesions on the cell membrane where the membrane contacts the extracellular
matrix. Along the actin microfilaments, myosin and its regulators linked with the lamellipodial
actin at the side of adhesion provide the motive force of the cell. The fourth and last stage of
cell migration is the release of adhesion in the rear end, allowing the cells to migrate in a
forward direction. The release of calcium ions that activate proteases that destroy the focal
adhesion sites from stretch-sensitive calcium channels is vital in this process.
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Source: https://www.researchgate.net/profile/Gary-Borisy-2/publication/8975413/figure/fig1/AS:280344570941448@1443850762676/Steps-in-cell-migration-Polarity-is-intrinsic-to-a-migrating-cell-A-Cdc42-along_Q640.jpg
Figure 14. Cell Migration
Induction is the kind of interaction of two or more cells or tissues of different histories
and properties at close range. The two components of inductive reactions are (1) inducer,
the tissue that produces the signal that changes the behavior of other tissues and (2) the
responder, the tissue being induced. It is highly significant that the responding tissues
should have both a receptor protein for the inducing factor and the ability to respond to the
signal. It is important to note that the presence of the receptor protein does not guarantee
that the tissue is competent to respond to the inducer’s signal. Furthermore, one induction
can give a tissue the competence to respond to another tissue. In contrast to endocrine
factors, such as hormones that travel through the blood, induction has paracrine factors
that alter the behavior or differentiation of adjacent cells.
There are two major modes of inductive interaction established by Howard Holtzer
(1968): (1) instructive interaction, where the signaling of an inducing cell initiates new gene
expression and (2) permissive interactions, where the responding tissue has already been
specified and requires only an environment that allows the expression of new traits.
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Examples of induction interactions occur between sheets of epithelial cells with

adjacent mesenchymal cell, called epithelial-mesenchymal interactions. These induction
interactions can be regional specificity or genetic specificity. Example of regional specificity
of induction is the formation of cutaneous structures, such as broad feathers of the wing,
narrow feathers of the thigh, or the scales and claws of the chick feet, by the secretion of
proteins from the chick epidermis to the underlying dermal cells to condense, which in turn
responds by secreting factors to the epidermis to form the structures. In genetic specificity of
induction, the experiment of Spemann and Schotte (1932) of the manipulation of the oral
structures of salamanders and frogs showed that mesenchymal tissue that instructs certain
formation of anatomical structures can cross species barriers but will only create the
anatomical structure it knows.
Figure 15. Regional and Genetic Specificity of Induction (Gilbert and Baressi, 2016)
There are several events of induction interactions, with some of them are the
following: (1) juxtracrine interactions which occurs when cell membrane proteins on one
cell surface interact with the receptor proteins on adjacent cell surfaces, (2) paracrine
interactions, where proteins synthesized by one cell diffuse over small distances to induce
changes in neighboring cells, and (3) autocrine interactions, which occurs when the same
cells that secrete paracrine factors also respond to them.
Group of Paracrine Factors
Paracrine factors can be grouped into four major families based on their structures:
(1) fibroblast growth factor (FGF) family, (2) hedgehog family, (3) the Wnt Family and
(4) the TGF-β superfamily.
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The fibroblast growth factor (FGF) is expressed from FGF genes, and is vital in
regeneration (Fgf1 protein), blood vessel formation (Fgf2), skin development (Fgf7), and limb
development and lens induction (Fgf8). Pathways where FGF inductions are involved is the
(1) RTK pathway, and (2) JAK-STAT pathway, which is essential in the differentiation of
blood cells, limb growth, and casein activation in milk production.
The hedgehog family are often utilized to induce certain cell types in an embryo and
to create boundaries between tissues. Vertebrates has three homologues of the Drosophila
hedgehog genes: (1) sonic hedgehog (shh), (2) desert hedgehog (dhh), and (3) Indian
hedgehog (ihh). The sonic hedgehog is responsible for ensuring that motor neurons arise
only from the ventral portion of the neural tube, that portion of each somite forms the
vertebrae, the proper places of the growth of feathers, and that the pinkies are always the
most posterior digits of vertebrate hands and feet. The desert hedgehog is important in
spermatogenesis in mice, while Indian hedgehog is expressed in the gut and cartilage and
is essential in postnatal bone growth. The proteins of the Hedgehog family is involved in the
Hedgehog Pathway necessary in vertebrate limb development, neural differentiations and
facial morphogenesis.
The Wnt family are vital in establishing the polarity of vertebrate and insect limbs,
stem cell proliferation, and the development of urogenital system development. The proteins
in this family is involved in the ‘canonical’ and ‘non-canonical’ Wnt pathways. The TGF-β
superfamily are vital in the formation of the extracellular matrix between cells and for the
regulation of cell division (TGF-β1, 2, 3, and 5). Included in this superfamily is the bone
morphogenetic proteins (BMP) that induce bone formation, regulate cell division, apoptosis,
cell migration and differentiation. The SMAD Pathway is also activated by the members of
this superfamily.
Other paracrine factors include epidermal growth factor, hepatocyte growth factor,
neurotrophins, and stem cell factors. They do not belong to the other groups because of
differences in structures. Some paracrine factors are involved exclusively on some
developments such as the erythropoietin, cytokines and interleukins that are associated with
developing blood cells.
Juxtacrine Signaling
In juxtacrine signaling, the proteins from the inducing cell interact with receptor
proteins of adjacent, responding cells without diffusing. There are two widely used families
of juxtacrine factors used in this type of signaling: (1) Notch proteins and the (2) the eph
receptors and their ephrin ligands. The Notch proteins and its pathways are involved in the
formation of several organs such as the kidney, pancreas, and heart, as well as the receptors
in the nervous systems. They are also vital to the patterning of the nematode vulva.
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References
Gilbert, S. and Barresi, M.J.F (2016). Developmental Biology. Eleventh Edition. USA: Sinauer
Associates Inc.
Self Help:
Davies, J. A. (2005). Mechanisms of morphogenesis. ProQuest Ebook

Central https://ebookcentral.proquest.com.
Organ development. (2019). ProQuest Ebook Central https://ebookcentral.proquest.com
Let’s Check!
Activity 3. Now, that you know the basic of cell adhesion, cell migration and cell signaling, let
us try to check your understanding of these topics. Match the following items that is
associated with different events in cell-to-cell communication:
____1. Juxtacrine signaling a. Cell adhesion
____2. Polarization b. Cell migration
____3. Epithelial-mesenchymal interaction c. Cell signaling
____4. E- and P-cadherin
____5. Positive affinity of mesoderm
____6. Cell sorting
____7. Paracrine factors
____8. Lamellipodia
____9. Autocrine interactions
____10. Release of the rear end
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Let’s Analyze
Activity 3. Now that you are acquainted with cell adhesion, cell migration and cell signaling,
it is important that you will be able to relate this in your future profession. I will require you to
give what are asked:
1. Draw and discuss an example of cell death pathway. What is the significance of this
process for a certain organism?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. What are the different malformations caused by mutations in the Hedgehog genes?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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In a Nutshell…
Activity 3. Based on your learnings about the cell adhesion, cell migration, and cell signaling,
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
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Keyword Index
Cell-to-cell communication Induction
Cell Adhesion Cadherin
Cell Migration Ligands
Paracrine Factors Receptor
Juxtracrine Signaling Morphogenesis

ULO-b. Discuss the process of maintaining the differentiated state, and the role
of extracellular matrix, and the epithelial-mesenchymal transition.
Metalanguage

demonstrate ULO-b will be operationally defined to establish a common frame of reference
1. Extracellular matrix is an insoluble network of macromolecules secreted by cells

into their immediate surrounding.
2. Integrins belong to the family of proteins that attaches extracellular matrix and the
cells.
3. Anoikis is a type of programmed cell death that occurs in anchorage-dependent cells
when they detach from the surrounding extracellular matrix.
4. Epithelial-mesenchymal transition consists of orderly series of events where
epithelial cells are transformed into mesenchymal cells.
Key Knowledge
Maintaining the Differentiated State
To finalize the commitment of cells to become a particular specialized cell, it is

necessary that gene expression must be maintained. There are four pathways established
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by the process of evolution that aim to maintain the differentiation once gene expression is
initiated:
1. Binding of the transcription factor of genes activated by the signal transduction

cascade to the enhancer of its own gene. Through this, the transcription factor
will be made, and its synthesis will become independent of the signal that induced
it originally.
2. Maintaining gene accessibility by synthesizing proteins that act on chromatin.
3. Synthesis of signaling molecule and molecule receptor by the induced cell if the
differentiation is dependent on other cell signaling. Autocrine factors can sustain
the induction and complete their differentiation after cell induction.
4. Interaction of cells to with its neighboring cells such that each one stimulates the
differentiation of the other, and the part of each neighbor’s differentiated
phenotype is the production of a paracrine factor that stimulates the other’s
phenotype.
The Extracellular Matrix
Cell adhesion, cell migration and cell signaling greatly depend on the extracellular
matrix. The extracellular matrix is a network of macromolecules secreted by cells in their
surrounding environment. It has collagen, proteoglycans, and specialized glycoproteins
molecules such as the fibronectin and laminin. It serves as a substrate in which the cells can
adhere, migrate, and communicate. Proteoglycans aids in the delivery of paracrine factors.
Examples are heparan sulfate, that bind many members of the TGF-β, Wnt and FGF
families, and chondroitin sulfate proteoglycans. Fibronectin serves as general adhesive
molecule that links cells to one another and other substrates such as collagen and
proteoglycans, proper alignment of cells, and paves a road for migrating cells. Laminin and
type IV collagen make up the basal lamina, the extracellular matrix that surrounds epithelial
tissues. Laminin aids in the assembly of the extracellular matrix, thus contributing to cell
adhesion and growth, cell shape, and cell migration.
The receptors for the extracellular matrix molecules are the integrins. Integrins
integrate the extracellular and intracellular scaffolds, allowing them to work together. An
example of integrin is the receptor in the fibronectin receptor complex, in which it binds the
fibronectin of the extracellular matrix, and on the inside of the cell, it serves as an anchorage
site for the actin microfilaments that move the cell. On the extracellular side, integrins bind to
the sequence arginine-glycine-aspartate found in extracellular matrices such as fibronectin,
vitronectin and laminin. In the cytoplasmic side of the cell, the integrins bind to talin and α-
actinin, which bind to actin microfilaments, allowing the cell to move during migration.
Inducing specific gene expression can also be done by extracellular matrix such as
in liver, testis and mammary glands. For instance, the chondrocytes that produces the
cartilage of the vertebrate and limbs can survive and differentiate only if they are surrounded
by extracellular matrix and joined to it by integrins. If these cells are not able to attach
themselves, they die causing a special type of apoptosis called anoikis.
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The Epithelial-Mesenchymal Transition
The Epithelial-Mesenchymal Transition (EMT) consists of orderly series of events

where epithelial cells are transformed into mesenchymal cells. It provides a summary that
integrates the processes of cell-to-cell communication. It is very essential during
development especially during the formation of neural crest cells from the dorsal most region
of the neural tube, formation of mesoderm and the formation of vertebrae precursor cells
from the somites.
During EMT, normal epithelial cells are attached to one another through adherens
junctions containing, cadherins, catenins and actin rings. They are attached to the basement
membrane through the integrins. Expression of genes that encode for these cellular
components is repressed by the paracrine factors, causing the cell to lose polarity, lose
attachment to the basement membrane and cohesion with other epithelial cells. Because of
this repression, cytoskeletal remodeling occurs, as well as the secretion of proteases and
extracellular matrix molecules that enable the migration of the newly formed mesenchymal
cells.
Figure 15. Epithelial-Mesenchymal Transition (Gilbert and Baressi, 2016)
References
Associates Inc.
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Self Help:
Davies, J. A. (2005). Mechanisms of morphogenesis. ProQuest Ebook

Organ development. (2019). ProQuest Ebook Central https://ebookcentral.proquest.com
Let’s Check!
Activity 4. Now, that you know the basics of maintaining the differentiated state, extracellular
matrix and the epithelial-mesenchymal transition, let us try to check your understanding of
these topics. Give what is asked:
1. Draw a figure of an integrin-in-action. Discuss briefly.
________________________________________________________________________
________________________________________________________________________
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________________________________________________________________________
________________________________________________________________________
Let’s Analyze
Activity 4. Now that you are acquainted the basics of maintaining the differentiated state,
extracellular matrix and the epithelial-mesenchymal transition, it is important that you will be
able to relate this in your future profession. I will require you to give what are asked:
1. What is the role of anoikis in cancer progression?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Maintaining the differentiated state of cells is important in the process of development.

Give examples of cells that maintain their differentiated form through:
a. Binding of transcription factors to its own enhancer
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
b. Maintaining gene accessibility

_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
c. Synthesizing of own inducing and receptor molecule

_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
d. Differentiation through interactions between neighboring cells

_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
_______________________________________________________________
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In a Nutshell…
Activity 4. Based on your learnings about the basics of maintaining the differentiated state,
extracellular matrix and the epithelial-mesenchymal transition, feel free to write your
arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
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5.
Keyword Index
Enhancer Extracellular matrix
Epithelial-Mesenchymal Transition Integrins
Anoikis Ligands
Paracrine Factors Proteoglycans
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Big Picture
FERTILIZATION AND EARLY DEVELOPMENT IN VERTEBRATES

a. Discuss the concepts of fertilization specifically gamete structure, and external and
internal fertilization in invertebrates and mammals, respectively.
b. Discuss the early development in invertebrates such as in sea urchins, snails,
tunicates, nematodes and fruit flies.

ULO-a. Discuss the concepts of fertilization specifically gamete structure, and
external and internal fertilization in invertebrates and mammals, respectively.
Metalanguage

1. Fertilization involves the fusion of opposite gametes to create a new

individual with different genome derived from both parents.
2. Gametes are the mature reproductive cells involved in fertilization.
3. Capacitation is the maturation of sperm within the female reproductive tract.
4. Oocyte is a cell in the ovary that undergoes meiotic division to form an egg
or ovum.
5. Ovum, or egg, is the immobile female gamete.
6. Pronuclei is either of the pair of the gamete during fertilization but before
their fusion leads to the formation of the diploid nucleus of the zygote.
Key Knowledge
Generally, fertilization involves the fusion of sperm and egg. Through fertilization, a
new set of genomes derived from two parents is created in a new offspring, thus, encouraging
the diversity of life. In addition, fertilization also initiates further cell development. In general,
fertilization consists of four (4) major events: (1) contact and recognition between the egg
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and the sperm, (2) sperm entry regulation into the egg, (3) fusion of the genetic
material of the egg and the sperm, and (4) activation of egg metabolism to start
development.
Structure of the Gametes
Sperm Structure
Generally, the sperm cells across species consist of a haploid nucleus, a sac of
enzymes for the entry of nucleus into the egg, and a propulsion system that moves the
sperm cells after ejaculation. The head of the sperm consist of the acrosome and the nucleus.
The acrosomal vesicle, or acrosome, is a modified secretory vesicle derived from the Golgi
apparatus that digests proteins and complex sugars, paving the way of the sperm’s nucleus
into the cell. Acrosomal process is a structure that allows recognition between the egg and
the sperm. This is useful especially to animals that have external fertilization, such as in sea
urchins.
The propulsion system of the sperm consists of the flagellum. Generally, the sperm
has a major motor protein called the axoneme, and dynein, which provides the force for
sperm propulsion. In most mammals, the sperm becomes completely mature when it has
been inside the female reproductive tract for a period of time. This is called capacitation.
Source: https://s3-us-west-2.amazonaws.com/courses-images/wp-content/uploads/sites/1940/2017/06/23191325/1024px-Simplified_spermatozoon_diagram.svg_-1024x298.png
Figure 16. Sperm Structure
Egg Structure
Typically, the egg, or the ovum, contains the materials necessary for the early stages
of development. During the development of the ovum, the oocyte, the cell in the ovary before
it undergoes meiosis, accumulates more material in its cytoplasm in preparation for
fertilization and afterwards, growth and development. When the egg already matures, its
cytoplasm now contains: (1) nutritive proteins, (2) ribosomes and tRNA for the synthesis
of structural proteins and enzymes by the embryo, (3) messenger RNA that contains
information for protein production in early development stages, (4) morphogenic factors
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that influence cell differentiation and (5) protective chemicals that protect the developing
immobile embryo.
The egg also contains a large nucleus. Female pronucleus, such as in sea urchin,
is already haploid at the time of fertilization. Some species, such as in some mammals and
worms, the egg nucleus becomes haploid through meiotic divisions only when the sperm, or
male pronucleus, enters the egg. The egg is covered in a cell membrane that plays a big
role in recognizing sperm and regulation of efflux and influx of ions during fertilization.
Extracellular matrix also surrounds the egg and it also helps in the recognition between
sperm and egg. In invertebrates this external structure is called the vitelline envelope. Some
species have an additional layer of egg jelly outside the vitelline envelope that attract or
activate sperm. In mammals, the egg has a separate extracellular envelope called the zona
pellucida and cumulus, layer of cells from the ovarian follicular cells that nurtures the egg
after it is released from the ovary. Corona radiata is the innermost layer of cumulus cells
adjacent to the zona pellucida.
Figure 17. Sea urchin egg cell surface (Gilbert and Baressi, 2016)
Figure 18. Mammalian egg before fertilization (A. hamster egg fertilization; B. mouse
oocyte) (Gilbert and Baressi, 2016)
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The cortex, a thin layer of stiffer gel-like cytoplasm, can be found lying beneath the
cell membrane. It contributes to the formation of microfilaments necessary during cell division.
In addition, it also aids in sperm entry into the cell. Furthermore, the cortex also contains
cortical granules that also contain proteolytic enzymes that is homologous to acrosomal
vesicles. The difference is that the cortex contains approximately 15,000 cortical granules
that prevent polyspermy, or the entry of an additional sperm after a sperm has entered
already.
External and Internal Fertilization
Generally, sperm and egg interact with each other during fertilization as follows: (1)
the attraction of the sperm to the egg by the chemical substances produced by the
latter, (2) the release of enzymes from acrosomal vesicle by exocytosis, (3) the binding
of the sperm to the vitelline envelope or to the zona pellucida of the egg, (4) the
passage of the sperm through the mentioned extracellular matrices, and (5) the fusion
of the sperm and egg cell membrane. This process still varies across organisms. For
instance, in mammals, the sperm binds to the extracellular matrix before releasing the
contents of the acrosomal vesicles.
External Fertilization in Sea Urchin
Among organisms, the sea urchin has become the model organism for observing
external fertilization. In nature, the sea urchin is known for releasing its gametes to the
immediate surrounding, depending on the science of chemotaxis for the gametes to locate
each other and undergo the process of fertilization. Specifically, this assurance of fertilization
is highly dependent on the mechanisms of species-specific sperm attraction and species-
specific activation.
Figure 19. Summary of events during external fertilization in sea urchins (Gilbert and
Baressi, 2016)
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Sperm Attraction. Physico-chemical conditions that initiate the movement of aquatic

sperms varies across species. For sea urchin, the sperm starts to be mobile when it is
released into the environment. During the release, the sperm pH is elevated to approximately
7.6, thus, activating the dynein ATPase that provides the energy for the flagella to wave and
swim towards the egg. For the sea urchin Arbacia punctulata, the sperm is attracted to the
amino acid peptide, resact, diffused by the egg from its jelly. In addition to attracting the
sperm, resact also activates the sperm.
Acrosome Reaction and Recognition of the Egg’s Extracellular Coat. After

locating the egg, the acrosomal process in the sperm initiates contact with the egg jelly, and
this triggers the exocytosis of the sperm’s acrosomal vesicle, thus, releasing digestive
enzymes, that digest a path through the jelly coat to the actual egg. According to studies, the
acrosome process is coated with proteasomes from the acrosomal vesicle, and with this, it
can digest the vitelline envelope at the point of attachment to have access to the actual egg.
It is necessary to note that this acrosome reaction, as well as the recognition of the sperm to
the egg’s extracellular coat, varies across sea urchin species, thus, providing a set of
species-specific recognition events. In sea urchins, the acrosomal protein that mediates this
recognition in sea urchins is called bindin. This protein also varies in their sequences across
sea urchin species.
Fusion of Egg and Sperm Cell Membrane and Blocking Polyspermy. Fusion of
the egg and sperm cell membrane occurs after the sperm travels to the egg and the
acrosome reaction has undergone. In sea urchins, the egg can fuse with the sperm in all
regions of its cell membrane. For some species, however, there are only certain regions
where fusion can occur, depending on specific fusogenic proteins present. Bindin can also
contribute to fusing the sperm and egg cell membrane. Once the fusion has occurred and
the sperm has entered the egg, it is necessary that polyspermy does not occur. In sea urchin,
polyspermy can be prevented by a fast reaction, involving an electric change in the egg cell
membrane and a slower reaction, caused by the actions of cortical granules. The fast
block to polyspermy involves providing a selective barrier within the egg cytoplasm and its
environment to change the ion gradient in the two sides, thus, preventing the entry of sperm
for a few seconds. The cortical granules involved in the slow block to polyspermy is
dependent on the efflux or influx of calcium ions within the cell or from the environment. In
sea urchin and in mammals, calcium ions used to initiate cortical granules exocytosis comes
from the egg cytoplasm (intracellular) and not from the influx of calcium ions to the egg from
the surrounding. The release of calcium ions is also essential in activating the egg’s
metabolism and initiating development by freeing maternal mRNAs from cellular inhibitors.
Furthermore, the components of the cortical granules will bind to the vitelline envelope to
form a fertilization envelope that starts to form at the site of sperm entry, continuing its
expansion around the egg. The formation of fertilization occurs about 20 second after sperm
attachment and is complete by the end of the first minute of fertilization.
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Figure 20. Cortical granule exocytosis and formation of the fertilization envelope in
sea urchins (Gilbert and Baressi, 2016)
Figure 21. Calcium release across sea urchin egg during fertilization (Gilbert and
Baressi)
Fusion of the Male and Female Pronucleus. As the sperm’s nucleus and centrioles
enter the egg, thus, leaving its mitochondria and flagellum behind, it undergoes a process
where it decondenses to form the haploid male pronucleus. Eventually, the sperm centriole
will extend its own microtubules, integrating the with egg microtubule to form an aster. The
microtubules will contact the female pronucleus, causing the two haploid pronuclei to migrate
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towards each other and forms the diploid zygote nucleus. The synthesis of DNA can either
begin during migration, or pronuclear stage, or after the fusion of the pronuclei and the
formation of the zygote nucleus. Further development is then highly dependent on the
diffusion and level of calcium ions released earlier in fertilization.
Internal Fertilization in Mammals
Translocation and Capacitation of Gametes. In mammals, the translocation and

maturation of both sperm and egg cell are completed within the female reproductive tract.
This is achieved by the combination of small-scale biochemical interactions and large-scale
physical propulsion to arrive at the region of the oviduct where fertilization takes place called
the ampulla. The oviduct is commonly called as the uterine tube or the fallopian tube. In
translocation, the sperm gets into the oviduct by the muscular activity of the uterus. On the
other hand, a combination of ciliary beating and muscle contractions translocate the female
oocyte-cumulus complex from the ovary to its appropriate position for fertilization in the
oviduct.
In capacitation of the sperm, there are five sets of molecular changes observed: (1)
the sperm cell membrane is altered by the removal of cholesterol by the albumin present in
the female reproductive tract, (2) certain proteins and carbohydrates which may hinder
recognition between the sperm and the zona pellucida are lost, (3) enhancing the entry of
calcium ions into the sperm by altering the membrane of the sperm cell membrane potential
to negative, (4) protein phosphorylation to aid in the binding with the zona pellucida, and
lastly, (5) the outer acrosomal membrane is altered and travels to have contact with the
sperm cell membrane to prepare for fusion. It was observed that sperms that arrive early on
the oviduct has lesser chance of fertilizing the egg due to incomplete capacitation. It was also
observed that incapacitated sperm tend to bind actively with oviduct cells to increase its life
span and slow down the process of capacitation which may contribute to increasing the
probability that the sperm will still be able to meet the egg in the ampulla. This binding action
can also be an early block to polyspermy in a different perspective.
During capacitation, sperms are observed to be hyperactivated, that is, they swim at
higher velocities and generate greater force. These hyperactivated sperms can release
themselves from binding on the oviduct and continue their journey to the egg. In addition to
hyperactivation, it was also observed the sperm has hyaluronidase enzymes on the outside
of its cell membrane, enabling the sperm to digest a path through the cumulus cells. For the
sperm to locate the egg, a temperature gradient is formed within the oviduct. In second to
this heat sensing mechanism of the sperm is chemotaxis, where the egg along with the
cumulus cells secrete molecules that attract the sperm towards the egg during the last stages
of egg migration. Human follicular fluid is said to be involved during this chemotactic activity.
In addition, chemotaxis is generally attributed to the hormone progesterone produced by
the cumulus cells. Both physical and chemical sensing mechanisms are often seen in
capacitated sperms only.
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Binding of the Sperm to the Egg’s Zona Pellucida. The binding of the sperm to
the egg’s zona pellucida is relatively, but not absolutely species specific and varies across
species. In mouse egg, the zona pellucida has major glycoproteins (ZP1, ZP2 and ZP3) that
binds to the sperm to initiate acrosome reaction. The sperm contacts the egg at its side, in
contrast to sea urchin, where the sperm contacts the egg at its tip. After the acrosome
reaction, the contents of the acrosomal vesicle is released and a portion of the inner
acrosomal membrane is exposed. The junction between the inner acrosomal membrane and
the sperm cell membrane called the equatorial region becomes the site of membrane fusion
between the sperm and the egg. The sperm is then headed to the regions of the egg where
microvilli from actin polymerization extends to the sperm. In mammals, slow block to
polyspermy is initiated through the release of enzymes from the cortical granules.
Fusion of the Male and Female Pronucleus. The fusion of genetic material in
mammals takes longer than in sea urchin (12 hours). The sperm approaches the surface of
the egg tangentially, fusing with numerous microvilli. The DNA of the sperm, bounded in the
sperm chromatin by proteins called protamines, is uncoiled by glutathione in egg cytoplasm.
The oocyte is still at its metaphase of its second meiotic division when the sperm enters. The
entry of the sperm inactivates MAP kinase, allowing DNA synthesis, and other kinases that
leads to the proteolysis of cyclin, allowing cell cycle to continue and the formation of the
haploid female pronucleus, and securin, releasing the metaphase chromosomes at the
metaphase plate. Furthermore, calcium ion release plays a huge role in these events. With
this DNA synthesis continues in both male and female pronuclei, and the centrioles
accompanying the male pronuclei create asters from proteins of the oocyte. The microtubules
formed allows the two pronuclei to migrate and join one another. The two nuclear envelopes
break down upon meeting and the chromatins from both pronuclei condense into
chromosomes that orient themselves in a common mitotic spindle. Along with the sperm’s
male pronuclei and centriole is its mitochondria, and a small amount of cytoplasm, however,
the mitochondria and their DNA are degraded in the egg cytoplasm, thus, all the mitochondria
of the new individual is from the mother.
Activation of the Mammalian Egg. Lastly, the mammalian egg is activated by

waves of calcium ions triggered by the sperm. In contrast to sea urchin eggs that are
activated by a single wave of calcium ions, mammalian eggs are activated by numerous
calcium ion waves. This event can last for several hours and ends when egg activation is
complete. Egg activation is said to be complete when cortical granule exocytosis occurs, the
meiosis continues, and the maternal mRNAs are released from the inhibitors, respectively.
References
Associates Inc.
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Self Help.
The zebrafish: Cellular and developmental biology, part a cellular biology. (2016).
Kloc, M., & Kubiak, J. Z. (2014). Xenopus development. ProQuest Ebook
Let’s Check!
Activity 5. Now, that you know the processes of both external and internal fertilization, let us
try to check your understanding of these topics. Match the events and the stages of either
external or internal fertilization. Write your answers in the space provided:
_____ 1. The release of maternal mRNAs from inhibitors a. External Fertilization
_____ 2. Egg activation by calcium ion activities b. Internal Fertilization

_____ 3. Chemotaxis by the hormone progesterone c. Both
_____ 4. Cortical granules exocytosis
_____ 5. Capacitation
_____ 6. Sperm attraction through the protein resact

_____ 7. Formation of fertilization envelope
_____ 8. Sperm binding to the zona pellucida
_____ 9. Nurturing of oocyte by the cumulus cells
_____ 10. Fast block of polyspermy though changes in the electric charge of the egg cell
membrane
Let’s Analyze
Activity 5. Now that you are acquainted the processes of both external and internal
fertilization, it is important that you will be able to relate this in your future profession. I will
require you to give what are asked:
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1. How does the proteins axoneme and dynein function in sperm cells?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. What are the challenges encountered when studying mammalian fertilization? What
are the methods that aim to bridge these gaps?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. What are the differences between pronuclei fusion of the gametes during internal and
external fertilization?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. Some animals of different species, such as lions and tigers, can interbreed and
produce an offspring (a hybrid). How is this possible? Will the offspring be viable for
reproduction?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. In humans, how do birth control methods such as contraceptive pills, hormone IUD,
birth control (depo) shots and hormonal implants prevent conception?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 5. Based on your learnings about the basics of maintaining the differentiated state,
extracellular matrix and the epithelial-mesenchymal transition, feel free to write your
arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
Keyword Index
Fertilization Acrosome Process
Cortical Granules Polyspermy
Fertilization Envelope Pronucleus
Zona Pellucida Capacitation
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ULO-b Discuss the early development in invertebrates such as in sea urchins,
snails, tunicates, nematodes and fruit flies.
Metalanguage

1. Cleavage furrow is the indentation of the cell’s surface that start the progression
of cleavage.
2. Autonomous specification is specification that does not depend on the location
of the cell, rather, it depends on the cytoplasmic content of the cell.
3. Conditional specification is specification that depends on the location of cells,
and communication of cells with neighboring cells.
Key Knowledge
Cleavage
After fertilization, cleavage, or the rapid mitotic cell division of the fertilized cells into
smaller cells called blastomeres. This process is achieved by two coordinated process –
karyokinesis and cytokinesis. Karyokinesis is the mitotic divisions of the cell’s nucleus and
its mechanical agent is the mitotic spindle with tubulin microtubules, while cytokinesis
is the actual division of the cell with contractile ring of microfilaments made of actin as
its mechanical agent. In invertebrates, this process occurs rapidly to restore the somatic ratio
of nuclear volume to cytoplasmic volume. Growth of the zygote does not occur by the cell
skipping the gap periods of the cell cycle which is the G1 and G2 phase.
There are several patterns of embryonic cleavage. It is determined by two factors:

the distribution and amount of yolk protein within the cytoplasm, and the factors in the egg
cytoplasm that may influence the mitotic spindle’s angle and the timing of its formation.
Organisms that have eggs with sparse, equally distributed yolk are considered as isolecithal
and their cleavage is holoblastic, which means that the cleavage furrow extends through
the entire egg. In holoblastic cleavage, four major patterns are further observed: radial,
spiral, bilateral, and rotational. Examples of organisms with holoblastic eggs are sea
urchins, snails, and mammals. Those organisms with large amount of yolk in their eggs have
meroblastic cleavage, which means that the cleavage furrow does not penetrate the yolky
region of the cytoplasm. Centrolecithal eggs, like insect eggs, have yolk in the center with
the division of the cytoplasm occur in the rim of the cytoplasm. Telocithal eggs, like fish and
bird eggs, have a small portion of the area of the egg free of yolk. This area is where cell
divisions occur.
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Figure 22. Patterns of Cleavage (Gilbert, 2010)
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Gastrulation
Gastrulation occurs after cleavage. There are five (5) basic cell movements during
gastrulation: (1) invagination, where a region of cell in folds, (2) involution, or the inward
movement of an expanding outer layer, so that it spreads over the internal surface of the
remaining external cells, (3) ingression, or the migration of the individual cells from the
surface layer into the embryo’s interior, making the cells mesenchymal, (4) delamination, or
the splitting of one cellular sheet into two more or less parallel sheets, and (5) epiboly, or
the movement of epithelial sheets that spread as a unit to enclose the deeper layer of the
embryo.
Source: https://ib.bioninja.com.au/_Media/gastrulation_med.jpeg
Figure 23. Cell movement during gastrulation
Cell specification is achieved through cell-cell signaling. In addition, the asymmetric

distribution of positioning of patterning molecules during cleavage also influence cell fates.
There are three mechanisms to this patterning: (1) the molecules are bound to the egg
cytoskeleton and are acquired by the cells that obtain the cytoplasm, (2) the molecules are
transported along the cytoskeleton of other cells actively, and (3) the molecules can be
associated with a specific centromere that may follow one of the mitotic sister cells during
mitosis. These processes establish the asymmetry of the organism and in consequence,
specification between neighboring cell will be established through paracrine or juxtacrine
interactions. The three axes that serve as the foundation for the embryo’s body – anterior-
posterior axis, dorsal-ventral axis, and left-right axis – should also be determined during this
span of the development. In this section, the embryogenesis of four invertebrates will be
briefly explored. According to Davidson (2001), this embryogenesis is categorized as “TypeI
embryogenesis’, which includes the immediate activation of the zygotic genes, rapid
specification of the blastomeres by the products of the zygotic genes and maternally active
genes, and a relatively small number of cells at the start of gastrulation.
Early Development in Sea urchin
Sea urchin cleavage exhibit radial holoblastic cleavage. Based on studies, most of
the embryonic cell fate are already specified during the 60-cell stage, however, the cells are
not irreversibly committed, which means that the cells remain pluripotent and can still rise to
other cell types. Sea urchin cell fates are determined by both conditional and autonomous
specification. The patterns of cell division during cleavage becomes less regular when the
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embryo becomes a 120-cell blastula. A blastula is a hollow sphere with cells surrounding a
central cavity or blastocoel. During this stage, all the cells are of the same size. Even though
division still occurs, the blastula remains to be one-cell layer thick. By the ninth and tenth
division, the fates of the cells have already become specified, with each cell becoming ciliated
on the region of the cell membrane located in the farthest part from the blastocoel. The cilia
will accommodate the embryo’s mobility later. The cells in the vegetal pole will begin to
thicken and form a vegetal plate, while the cells of the animal hemisphere will create hatching
enzymes that will digest the fertilization envelope. When the embryo hatches from the
envelope, it will become a free-swimming hatched blastula. The anterior-posterior axis, which
is influenced by the animal-vegetal axis, is already established prior to fertilization. Both
dorsal-ventral and left-right axes are specified after fertilization.
Source: https://lh3.googleusercontent.com/proxy/qvkq_MKDhbanZDAC1wE6vELRvcfaTE1cIiOKjeqYzLgwa-
BTkuKGMlRFN5Kd1PGLuCltY8Zz0sLNJ7ipBfasxAKfDTRFRCuBtPA05QRlPL8arZwuR50ccgpYuA
Figure 24. Sea Urchin Cleavage
Sea urchin gastrulation consists of several two (2) key events: (1) ingression of the
skeletogenic mesenchyme, and (2) invagination of the archenteron. Skeletogenic
mesenchyme, also called primary mesenchyme, forms the larval skeleton. The cells that
migrate for the ingression of the skeletogenic mesenchyme are from the small blastula cells
called micromeres located at the farthest end of the embryo’s vegetal pole, specifically, the
large micromeres. These large micromeres enter the blastocoel through cell migration by
using filopodia and become localized in the ventrolateral region of the blastocoel. They will
fuse into syncytial cables that will serve as the axis of the calcium carbonate spicules of the
larval skeletal rods.
While the large macromeres are migrating, the specified cells remaining in the vegetal
pole, flattens and thickens to form a vegetal plate. In addition, these cells also fill thegap
caused by the migrating large micromeres. The vegetal plate, then, involutes inward by
altering its shape, and then invaginates by a quarter to a half the way into the blastocoel
before stopping. The primitive gut, or the archenteron is created. The opening of the
archenteron at the vegetal pole is called blastopore. The primary cells that invaginate are
the non-skeletogenic mesenchyme, which will later form the musculature around the gut
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and the coelomic pouches. The endodermal cells adjacent to the micromere-derived
mesenchyme migrate the farthest distance into the blastocoel and becomes the foregut. The
midgut is formed by the next layer of endodermal cells and the last circumferential row that
invaginates from the hindgut and the anus. Further stages of archenteron invagination
Figure 25. Sea Urchin Gastrulation (Urry et al., 2017)
Early Development in Snails
Snails, along with some annelid worms, flatworms, and most mollusk, undergo
spiral holoblastic cleavage. Snail shell coiling is influenced by the orientation of cleavage
plane. Simultaneously, the orientation of cleavage plane is also controlled by cytoplasmic
factors in the oocyte. In some snail species, this direction is uniform, with some mutants
showing opposite coiling. Coils with opening to the left is called sinistral coiling while the
opposite is called dextral opening. In contrast to sea urchin’s development, snail have a
mosaic development, in which the blastomeres are specified autonomously. The mRNAs
and paracrine factors used during development is cytoplasmic localized in the specific region
of the oocyte. Snail gastrulation is accomplished through epiboly, where micromeres located
at the animal cap multiply and overgrow the vegetal macromeres. These micromeres will
cover the entire embryo and a small blastospore slit will be left at the vegetal pole. Since
snails are protostomes, the blastospore will develop into the snail’s mouth regions.
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Figure 26. Origin of Sinistral and Dextral Coiling Determined by the Direction of
theMitotic Spindle at the Second Cleavage (Gilbert, 2010)
Early Development in Tunicates
Tunicates are marine organisms that belong to the Phylum Chordata. At some point
of their life, they possessed a notochord and a dorsal nerve cord. As they undergo
metamorphosis, these two structures degenerate. Tunicates have bilateral holoblastic
cleavage. The cell division during cleavage is always asymmetrical, such that the anterior
cells are larger that the posterior blastomeres. Specification of tunicate blastomeres occur
both autonomously and conditionally. Organs that are developed through conditional
specification in tunicates include the brain, notochord, heart, and mesenchyme cells.
The axes of the tunicate larva are already determined by the cytoplasm of the zygote
before the start of the first cleavage. The dorsal-ventral axis is defined by the cap of
cytoplasm at the vegetal pole. The anterior-posterior axis is determined during fertilization,
where the migration of the oocyte cytoplasm occurs. Lastly, the left-right axis in tunicates is
determined during the first cleavage, however, the mechanism is poorly understood.
Gastrulation in tunicates is summarized by the invagination of the endoderm, the

involution of the mesoderm, and lastly, the epiboly of the ectoderm. Since they are
deuterostomes, the blastospore becomes the anus.
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Figure 27. Bilateral Symmetry in the egg of the tunicate Styela partita (Gilbert and
Baressi, 2016)
Early Development in Nematodes
In the nematode species, Caenorhabditis elegans, the zygote undergoes rotational

holoblastic cleavage. During cleavage, the axes of the nematode is already determined.
The anterior-posterior axis of the nematode is portrayed by the elongated axis of the
nematode’s egg. The dorsal-ventral and right left axes are seen in the division of the AB cell.
The AB cell, anterior founder cell, is the one of the two cells created by the first cell division
of the zygote. The other cell is called posterior stem cell (P1). Both conditional and
autonomous specification is seen in nematode embryo. Autonomous specification is evident
in the P1 lineages and conditional specification can be seen on the development of the
endoderm lineages. Gastrulation in C. elegans start early by the generation of the P4 cell (a
P1 lineage) in the 24-cell embryo. By 6-hours from fertilization, the embryo cells would have
developed into organs, and the ball-shaped embryo stretches out to become a worm.
Source: https://www.mun.ca/biology/desmid/brian/BIOL3530/DEVO_06/ch06f01.jpg
Figure 28. Early Embryonic Development in Nematode
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Early Development in Drosophila
In the genus Drosophila (fruit flies), egg activation occurs before the sperm enters
the egg. It is accomplished during ovulation, a few minutes before fertilization begins. In
addition, axes specification also occurs before the sperm meets the egg. The sperm can only
enter at one site called the micropyle which is located at the future dorsal anterior region of
the embryo. Unlike newly fertilized sea urchin and mammalian cells, there is no cortical
granules in Drosophila zygote that can act against polyspermy. Instead, the micropyle is said
to be analogous to the function of cortical granules. Drosophila undergo superficial
cleavage. This type of cleavage is characterized by the confinement of cleavage to the
cytoplasmic rim of the egg because of the presence of a centrally located yolk. In addition,
cells do not multiply unless the nuclei have divided several times, creating a syncytium – a
single egg with many egg in its cytoplasm. After ten cellular divisions, the zygote will be called
a syncytial blastoderm. During this stage, five nuclei will migrate towards the posterior pole
of the embryo, where they will be enclosed by cell membrane and become pole cells. These
pole cells will give rise to adult gametes.
As the cells is formed from the syncytial blastoderm, the Drosophila embryo will
undergo a mid-blastula transition. During this time, the cleavage becomes asynchronous
and new mRNA is created. During this time, maternal control of development will be
transferred to the zygote. The gastrulation of the Drosophila embryo starts with the
invagination of the most ventral region of the embryo, causing the formation of a ventral
furrow. The germ band expands such that the future posterior segments curl behind the
presumptive head. There are several genes involved in the next stages of Drosophila
development. These are summarized on the figure below.
Source: https://clinicalgate.com/wp-content/uploads/2015/06/B9781455727940000048_f004-001-9781455727940.jpg
Figure 29. Genes Involved in the Development of Drosophila Embryo
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References
Associates Inc.
Self Help.
Caenorhabditis elegans: Cell biology and physiology. (2012). ProQuest

EbookCentral https://ebookcentral.proquest.com
Let’s Check!
Activity 6. Now, that you know the basics about early development in some invertebrates,
let us try to check your understanding of these topics. Match the events of early
development to its corresponding invertebrate. Write your answers in the space
provided:
1. Mid-blastula transition a. Sea Urchin
2. Synctial blastoderm b. Snail
3. Autonomous specification c. Tunicate
4. Conditional specification d. Nematode
5. Mosaic development e. Fruit fly

6. Rotational holoblastic cleavage
7. Bilateral holoblastic cleavage
8. Spiral holoblastic cleavage
9. Formation of anus from blastospore
________10. Formation of mouth from blastospore
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Let’s Analyze!
Activity 6. Now that you are acquainted with the basics about early development in some
invertebrates, it is important that you will be able to relate this in your future profession. I will
1. What is the difference between conditional and autonomous development?
2. At what stage of early development in snail are the axes determined? Explain.
3. At what stage of early development in tunicate is bilateral symmetry fixed? Explain.
4. At what stage of early development in Drosophila are the axes determined? Explain.
5. What are the different genes involved in the segmentation of the Drosophila
body?Explain.
In a Nutshell
Activity 6. Based on your learnings about the basics about early development in some
invertebrates, feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
Question and Answer’s List

6.

1.
2.
3.
4.
5.
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Keyword Index
Mid-blastula Transition Mosaic development
Blastospore Cleavage
Autonomous specification Axis
Conditional specification Blastoderm
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Big Picture
EARLY DEVELOPMENT IN VERTEBRATES AND LATE EMBRYONIC
DEVELOPMENT

a. Explain the processes of early embryonic development in vertebrates, specifically, in

amphibians, fishes, birds, and mammals.
b. Discuss the processes of later embryonic development.

ULO-a. Explain the processes of early embryonic development in vertebrates,
specifically in amphibians, fishes, birds, and mammals.
Metalanguage

1. Mid-blastula transition, in frogs, involves the transition of the embryo

performing its own genetic transcription from relying on maternally deposited
mRNAs. This is also called maternal zygotic transition.
2. Cortical rotation, in frogs, is the cytoskeletal shifting of the embryo, where the
outer layer of the cell, the cortex (cortical cytoplasm), rotates to the inner
cytoplasm.
3. Discoidal cleavage involves the cleavage on the small portion region of the
cytoplasm without yolk (blastodisc) in fishes and birds.
4. Primitive streak, in birds, reptiles and mammals, is a structure that forms in the
blastula that will establish bilateral symmetry, determine gastrulation site and
formation of germ layer.
Key Knowledge
Early Amphibian Development
Fertilization. In frogs, fertilization can occur either externally or internally of the

female’s body. Generally, a frog’s sperm can enter the egg cell in the animal hemisphere of
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the amphibian embryo. The dorsal-ventral axis is determined immediately during the sperm’s
entry. Generally, most frog’s species embryo undergoes cortical rotation, where the cortical
cytoplasm rotates with respect to the internal cytoplasm. In a 1-cell embryo, the cortical
cytoplasm rotates 30°. Cortical rotation is organized by the sperm centriole through
rearranging the microtubules of the egg in parallel array in the vegetal cytoplasm.
Experiments that inhibit cortical rotation in frogs resulted to the complete gain of ventral
features but complete loss of dorsal features. The gray crescent, a band of inner gray
cytoplasm in the marginal region of the 1-cell embryo, will be the starting point of gastrulation.
It is located directly opposite of the sperm entry point.
Source: https://lh3.googleusercontent.com/proxy/lRvDfC9VobTHoBCnP2Ik_7I3YHb1dK9c2i4wswxzucyFtPBogxCPpQhqgATOZJplnCR5gPKs6C_8uV6-g-oJouGnMaFwcz2ujBk6J3gKmxo
Figure 24. Cleavage in a Frog Embryo
Cleavage. Most frog and salamander embryo exhibit holoblastic, radially

symmetrical cleavage. Because amphibian eggs contain more yolk in the vegetal
hemisphere, cleavage in this hemisphere is slow compared to the animal hemisphere. The
first cleavage bisects the gray crescent. Before the first cleavage is completed in the vegetal
hemisphere, the second cleavage at the animal pole already starts. When the frog embryo
has 16 to 64 cells, it becomes a morula. At the 128-cell, the blastocoel is evident and the
embryo is now called a blastula. Mid-blastula transition starts late in the 12th cell cycle,
preparing the embryo for gastrulation. During this transition, nuclear genes become activated
and transcribed in different cells. Furthermore, the gap phases become present and the
blastomeres become motile.
Source: https://www.macmillanhighered.com/BrainHoney/Resource/6716/digital_first_content/trunk/test/hillis2e/asset/img_ch38/c38_fig10.jpg
Figure 25. Gastrulation in a Frog Embryo

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Gastrulation. In early gastrulation in frog embryo, the blastospore forms by the

invagination of cells in the marginal zone into the embryo, closer to the animal pole. The
marginal cells migrate into the embryo because of the animal pole’s rapid cell division and
flattening. The animal hemisphere, then, undergoes epiboly. The marginal cells involute to
line the inner wall of the animal cell sheet, starting close to the blastospore dorsal lip and
head towards the ventral side of the embryo until the marginal zone cells line the roof and
inner walls of the embryo. The blastocoel will be displaced by the marginal zone cells and
the archenteron forms between the cells lining the roof and cells which have just migrated
into the cell. The animal cap sheet of cells continues to wrap around the external surface of
the embryo. By late gastrulation, most of the embryo will now be covered by the animal cap
cell sheet and the blastocoel has almost been completely displaced by the archenteron
expansion. A yolk plug is formed due to the inhibition of involution of vegetal and marginal
cells due to the animal cap cell sheet. It is absorbed by the embryo later. By the end of
gastrulation, the blastula becomes a gastrula. The marginal cells that lines the internal walls
of the embryo form the mesoderm, the yolk plug made up of vegetal cells becomes
internalized and forms the endoderm, and the archenteron with the walls lined by mesoderm
will continue to develop into the gut.
Early Development in Fishes
Fertilization and Cleavage. Zebrafish has been the common model organism for
observing early development in fish embryo. Most fertilization in fishes occur externally.
Fertilization in zebrafish starts to activate cytoplasmic movements in about 10 minutes.
Waves of calcium ions, initiated by fertilization, results to the squeezing of the non-yolky
cytoplasm into the animal pole of the egg converting the spherical egg into a pear-shaped
structure with an apical blastodisc.
Figure 26. Cleavage in Zebrafish Embryo (Gilbert, 2010)
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Cleavage in zebrafish occurs about 40 minutes after fertilization. Since fish eggs are
telolecithal (most of the egg is occupied by yolk), cleavage takes place on the apical
blastodisc, resulting to meroblastic cleavage. The embryo forms from the blastodisc so the
type of meroblastic cleavage is referred to as discoidal. By the first 12 cell divisions, a mound
of cells sits at the animal pole of a large yolk cell, constituting the blastoderm. Mid-blastula
transition in observed during the tenth cell division, where zygotic gene transcription begins,
cell division slows down, and cell movement becomes evident. During this time, three distinct
cell populations can be observed: (1) the yolk syncytial layer, or YSL, formed when the
cells at the vegetal edge of the blastoderm fuse with the underlying yolk cell, (2) the
enveloping layer, or EVL, formed by the most superficial cells from the blastoderm, forming
an epithelial sheet, only a single cell layer thick, and the (3) deep cells located between the
EVL and the YSL that will give rise to the embryo proper.
Gastrulation. Gastrulation in zebrafish begins with the epiboly of the blastoderm, a

single multilayer of cells, over the yolk. During this stage, the deep cells of the blastoderm
move outward to intercalate or insert with more superficial cells. The rearrangements
thoroughly stir the positions of the deep cells, resulting to the thinning of the blastoderm and
its spread across the yolk cells. The second major cell movement of the gastrulation process
is the involution of cell at or near the blastoderm margin. This results to the folding of the
blastoderm into two cellular layers, called the epiblast and the hypoblasts, within a ring
called the germ ring around its entire circumference, forming the endoderm and mesoderm
in an anterior-posterior sequence according to the time of involution. The last major cell
movement during gastrulation involved the intercalations of mediolateral cells in both
the epiblast and hypoblast. This mediates the convergence and extension movements
towards the dorsal side of the gastrula, resulting to the dispersion of cells that were initially
neighbors along the anterior-posterior axis of the embryo.
Source: https://www.mun.ca/biology/desmid/brian/BIOL3530/DB_03/fig3_11.jpg
Figure 27. Gastrulation of Zebrafish Embryo

Early Development in Birds
Fertilization and Cleavage. Domestic chickens are used to observe early

development in birds. The fertilization of the chicken egg occurs at the oviduct prior to the
secretion of the albumen and shell that serve to cover it. Chicken egg are telolecithal, with
the blastodisc sitting atop a large yolk. It undergoes discoidal meroblastic cleavage after
fertilization and only occurs at the blastodisc, creating a single layer of blastoderm. Further
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cleavage division divides the blastoderm to a tissue 5-6 cell layers thick linked by tight
junction. The subgerminal cavity forms between the blastoderm and the yolk of the eggs,
when blastoderm cells absorb water from the albumen and secrete the fluid between them
and the yolk. The embryo forms in the part of the blastoderm called the area pellucida. This
part is only one cell thick and is the remaining cells among the deep cells located at the
center of the blastoderm that shed and die. The area opaca constitute of cells located at the
peripheral ring of blastoderm cells that did not shed their deep cells. The marginal zone is
located between the area pellucida and area opaca and it contain cells that is significant in
cell fate determination.
Source: https://veteriankey.com/wp-content/uploads/2016/07/B9780702028991000249_f1.jpg
Figure 28. Cleavage in Chicken Embryo
Gastrulation. Gastrulation in most avian embryo begins with formation of the

epiblast and hypoblast. Epiblast is the layer of cells from the area pellucida that remain at
the outer surface of the blastoderm. On the other hand, the hypoblast (hypoblast islands,
primary hypoblast, polyinvagination island) are area pellucida cells that have
delaminated and migrated individually into the subgerminal cavity. Secondary hypoblast
or endoblast forms when the anterior migration beneath the surface of sheet of cells derived
from yolky cells at the posterior margin of the blastoderm pushes the hypoblast islands
anteriorly. The two-layered blastoderm is joined together at the marginal zone of the area
opaca, forming a space between called the blastocoel. In some amniotes, or vertebrates
that undergo embryonic development within an amnion, extraembryonic tissues, such as yolk
sac, are present. Avian embryo, as well as the three germ layers, is from the epiblast only,
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while the hypoblast forms the external membranes such as the yolk sac and the stalk linking
the yolk mass to the endodermal digestive tube. In addition, the hypoblast also secretes
chemical signals that aid in the migration of epiblast cells.
Source: https://lh3.googleusercontent.com/proxy/cPMoil4XHd24SbPe7MgVY-ZABVIGEQwuBllyMe6BxNkbEYxauo5i-O8I9KsIyPkOZdYnH13NuiTJS5u724kEBxjC5JchSbWpg8GQFlvrwTc
Figure 29. The Epiblast and Hypoblast of Chicken Embryo
Gastrulation. The primitive streak is the major structural characteristic of avian,

mammalian and reptilian gastrulation. It will form the blastospore lips of the amniote embryo.
It arises from the posterior edge of the area called the Koller’s sickle and the epiblast located
above it. The cells will continue to migrate, converging at the primitive streak, causing it to
elongate. These cell movements will continue and the primitive streak will narrow and
lengthen, forming the primitive groove, or the chick blastospore. Migration of cells
continues inwardly through the primitive groove and the Hensen’s node. Hensen’s node,
also called the primitive knot, is an enlarged group of cells located at the anterior portion of
the primitive streak in a developing gastrula. Eventually, the cells generated in the Hensen’s
node and passing into the gastrula migrate anteriorly to form the head structures and the
notochord. The cells moving over the sides of the primitive groove will from the mesoderm
and endoderm while hypoblast will be displaced by the spreading of the endoderm. While
the posterior portion of the embryo is still part of the primitive streak and colonizing the
mesoderm, the cells located at the anterior end of the embryo has already started
organogenesis.
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Figure 30. Formation and Elongation of the Primitive Streak in Chicken Embryo
Early Development in Mammals
Fertilization and Cleavage. Fertilization in mammals also occur at the oviduct,

specifically, in the ampulla, a region close to the ovary. Cleavage in mammals is considered
as rotational cleavage and occurs 12-24 hours after fertilization. Meanwhile, the cilia in the
oviduct push the slowly developing embryo towards the cleavage. The first cleavage occurs
during this journey. Another characteristic of mammalian cleavage is the asynchronous
division of cells and compaction. In mouse embryo, compaction occurs at the third cleavage
where blastomeres, which are loosely arranged during the 8-cell stage embryo, huddle
together and form a compact ball of cell. This compaction is stabilized by tight junctions that
form between the cells outside the ball, while inside cells form gap junctions that enable the
passage of small molecules and ions. The compacted 16-cell embryo is now called a morula.
The inside cells, now called the inner cell mass, will give rise to the embryo proper, while
the descendants of the external cells become the trophoblast. The trophoblast will form
embryonic portion of the placenta called the chorion, the structure that enables the fetus to
get nourishment and oxygen from the mother. In addition, the chorion also secretes
hormones that enable the mother’s uterus to retain the fetus and regulators of the immune
response so that the mother will not reject the embryo.
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Source: https://bastiani.biology.utah.edu/courses/3230/db%20lecture/lectures/Cleavage/MamOvCleav.jpg
Figure 31. Fertilization and Cleavage in Mammals
Through the process of cavitation, the morula will form a blastocoel from the
secretion of fluid by the trophoblast cells. The blastocoel is enlarged by the accumulation of
sodium ions through sodium pumps. This is stimulated by the oviduct cells which travels
together with the embryo toward the embryo. Due to the expansion of the blastocoel, the
inner cell mass is positioned to one side of the ring of trophoblast cells. The resulting blastula
is now called a blastocyst. As the dividing embryo travels toward the uterus, it is necessary
that the blastocyst hatches from the zona pellucida. After hatching from the zona pellucida,
the blastocyst makes direct contact with the endometrium of the uterus. This process is
mediated by the protein L-selectin of the trophoblast cells. In addition, integrins, P-
cadherin, and proteases like collagenase, stromelysin, and plasminogen activator
ensures the success of the embryo attachment to the uterus.
Gastrulation. Unlike its ancestors of reptile and birds, mammalian embryos do not
depend on the stored yolk within the egg. Instead, mammals have evolved and restructured
maternal anatomy that involves the development of fetal organ capable of absorbing
maternal nutrients. In mammals, this is called the chorion, and is derived from the
trophoblast cells supplemented with mesodermal cells from the inner cell mass. The chorion
forms the fetal portion of the placenta and can induce uterine cells to from the maternal
portion of the placenta called the decidua. Blood vessels that will provide the oxygen and
nutrients to the embryo will become abundant in the decidua.
Segregation of cells in the inner cell mass will form two layers: the hypoblast
(primitive endoderm or visceral endoderm) and the epiblast, forming a structure called the
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bilaminar germ disc. The extraembryonic endoderm forms from the delamination of
hypoblast cells from the inner cell mass to line the blastocoel cavity. The extraembryonic
endoderm forms the yolk sac. On the other hand, the epiblast will split to small clefts that
coalesce to separate the embryonic epiblast from the other epiblast the lines the amnionic
cavity. The amnionic cavity fills with secretion called the amnionic fluid, functioning as a shock
absorber and keeping the embryo from being dry. The embryonic epiblast contains all the
cell that will generate the actual embryo.
Gastrulation in mouse embryo begins at the posterior end if the embryo, where the
cells of the node (analogous or refers to the Hensen’s node) arise. Like in chicken, the
mammalian mesoderm and endoderm migrate through a primitive streak. The cells arising
from the node give rise to the notochord. The precursors for the ectoderm are located at the
anterior and lateral to the fully extended primitive streak, however, these cells can arise to
descendants in more than one germ layer, or to both extraembryonic and embryonic
derivatives.
References
Associates Inc.
Self Help.
Caenorhabditis elegans: Cell biology and physiology. (2012). ProQuest

EbookCentral https://ebookcentral.proquest.com
Let’s Check!
Activity 7. Now, that you know the processes of early development in amphibians, fishes,
birds, and mammals, let us try to check your understanding of these topics. Match the
events/stages/concepts of early development and its corresponding group of animals they
occur. Write your answers in the space provided:
_____ 1. Embryo depending to stored yolk a. Amphibians

_____ 2. Formation of the primitive streak b. Fishes
_____ 3. Presence of the Hensen’s node c. Birds
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_____ 4. Blastodisc d. Mammals

_____ 5. Cavitation
_____ 6. Compaction
_____ 7. Cortical Rotation

_____ 8. Radial cleavage
_____ 9. Meroblastic cleavage
_____ 10. Mid-blastula transition
Let’s Analyze!
Activity 7. Now that you are acquainted with the processes of early development in
amphibians, fishes, birds, and mammals, it is important that you will be able to relate this in
your future profession. I will require you to give what are asked:
1. What causes ectopic pregnancy?
__________________________________________________________________
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2. How are twins formed? In terms of early development, what are the differences
between fraternal and identical twins?
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3. Draw a chicken egg. Name the parts of the chicken egg and state its function.
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4. Which part of the mammalian embryo is used as source of embryonic stem cells?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. State how axes are determined in:

a. Amphibians
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b. Fishes
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c. Birds
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_______________________________________________________________
d. Mammals
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_______________________________________________________________
In a Nutshell…
Activity 7. Based on your learnings about the processes of early development in amphibians,
fishes, birds, and mammals, feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
Keyword Index
Fertilization Mid-blastula transition
Cleavage Cortical rotation
Gastrulation Invagination
Discoidal Cleavage Delamination
Epiboly Involution
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ULO-b. Discuss the processes of later embryonic development including
emergence of ectoderm, paraxial and intermediate mesoderm, and lateral plate
mesoderm and endoderm.
Metalanguage

1. Organogenesis is the formation of organs within an embryo.

2. Competence refers to the cell’s ability respond to a specific inductive signal.
3. Specification is the first stage of cell or tissue commitment to a certain fate,
during which the cell or tissue can differentiate autonomously when placed in an
environment that is neutral with respect to the developmental pathway. This cell
commitment is still reversible.
4. Commitment describes a state where the cell’s developmental fate is already
restricted. The cell during this stage, however, is still not yet displaying obvious
changes in cellular biochemistry and function.
5. Differentiation is the process in which an unspecialized cell becomes
specialized into one of the many cell types.
6. Precursor cells is a widely used term to denote any ancestral cell type of a cell
lineage. Progenitor cells are undifferentiated cells that have the capacity to
divide few times before becoming differentiated.
7. Stem cells are undifferentiated cell with the capacity differentiate into specialized
cells. Stem cell potency is the capacity of the stem cell to generate different
types of differentiated cells.
8. Totipotent cells are stem cells capable of forming every cell in the embryo, and
the trophoblast cells of the placenta. Pluripotent cells are stem cells that can
become all the cell types of the embryo except the trophoblast. Multipotent cells
are stem cells in which their commitment and differentiation is already limited to
some cells of the body. Unipotent cells are stem cells that can only regenerate
a specific type of cell.
9. Neurulation is the process during vertebrate organogenesis that form the neural
plate (rise to become the CNS), epidermis, and the neural crest (rise to become
the PNS).
10. Mesenchyme cells are multipotent stem cells isolated from different sources.
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Key Knowledge
Emergence of Ectoderm
The germ layer ectoderm becomes the origin of an organisms’ central nervous
system (CNS), peripheral nervous system (PNS) and the epidermis. This process by which
the three ectodermal regions are made is called neurulation. The embryo undergoing this
process is called a neurula.
Figure 32. Major Derivatives of the Ectoderm Germ Layer (Gilbert & Baressi, 2016)
Construction of the Central Nervous System

Formation of the Neural Plate. The construction of the CNS in a vertebrate
organism begins with the formation of the neural tube from the neural plate. Neural plate cells
are often characterized by the expression of the Sox family. This family of transcription family
specify cells to become the neural plate and inhibit the formation of epidermis and neural
crest from the ectoderm.
Neural plate is formed during vertebrate gastrulation. It is the presumptive neural
tissue induced by the notochord. The notochord, a rod that extends along the dorsal side of
the vertebrate embryo, is derived from the dorsal mesoderm. The ectodermal cells above the
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notochord will be induced by the mesodermal cells and other tissues to become the neural
plate. In birds and mammals, the neural plate is formed opposite to the primitive streak where
ectodermal cell thickens and flattens.
Neurulation. Neurulation has two principal modes: primary neurulation (anterior)
and secondary neurulation (posterior).
Figure 33. Primary Neurulation (Formation of the Neural Tube) in Chick Embryo
(Gilbert & Baressi, 2016)
During primary neurulation, the invagination of the neural plate cells into the body
in order to separate and form an underlying hollow tube occurs. Generally, primary
neurulation included four distinct, but overlapping stages: (1) the elongation and folding
of the neural plate, where the edges of the neural plate thickens and move upward to form
the neural tube and the U-shaped neural groove; (2) bending of the neural plate, where
formation of hinge regions that remains in contact with surrounding tissues is formed (medial
hinge point or MHP in birds and mammals); (3) convergence of the neural folds, where
two dorsolateral hinge points (DLHPs) are induced and anchored to the surface ectoderm;
and lastly (4), closure of the neural tube, where the neural folds adhere together at the
dorsal midline and closes to create the neural tube and the apex of the neural folds
delaminate to become the neural crest cells.
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Secondary neurulation occurs at the most posterior region of the embryo during
tailbud elongation. It involves the production of mesenchyme cells from the prospective
endoderm and ectoderm cells. These mesenchyme cells condense into a medullary cord
beneath the surface of the ectoderm, exhibiting a mesenchymal-epithelial transition. After
this stage, lumens or hollow spaces are formed in the central portion of the cord, which will
eventually coalesce to become a single central cavity.
Figure 34. Secondary Neurulation in 25-Somite Chick Embryo (Gilbert & Baressi,
2010)
Brain. The brain is formed from the differentiation of the neural tube anterior-
posterior axis. In the anterior region of the neural tube, the neural tube balloons into three
primary vesicles: the forebrain, (prosencephalon), midbrain (mesencephalon) and the
hindbrain (rhombocephalon). The figure below shows the three primary brain vesicles and
the adult derivatives formed by the wall and cavities of the brain with their function in humans.
Figure 35. Early Human Brain Development (Gilbert & Baressi, 2016)
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Neuron Differentiation. The neuroepithelial cells of the neural tube gives rise to the
three different types of cells: (1) ventricular (ependymal) cells, which lines the neural tube
and secrete the cerebrospinal fluid; (2) precursors of the neuron, and (3) precursor of the
glial cells that helps in nervous system construction, insulation of neurons, and storage of
memories.
Tissues Architecture of the CNS. The neurons of the CNS are organized into layers
(cortices) and clusters (nuclei). The original neural tube is composed of a one-layer thick,
rapidly dividing neural stem cells called the germinal neuroepithelium. From this layer are
three-zone pattern of layers that forms the tissues of the developing CNS: (1) the ventricular
zone, where the stem cells continue to divide and the migrating cells form a second layer
called the (2) mantle, or intermediate zone, around the original neural tube; and (3) the
marginal zone. The cells in the mantle zone (gray matter) differentiate into both glial and
neurons, forming connections and sending their axons away from the neural tube lumen,
thus, the marginal zone (white matter) is poor in neuronal cell bodies. Eventually, these three
regions are retained during the development of the spinal cord and the medulla, which is the
posterior region of the hindbrain.
Figure 36. Development of the Human Spinal Cord (Gilbert & Baressi, 2016)
Spinal Cord and Medulla Organization. During the formation of the human spinal
cord, the neural tube is functionally divided into dorsal and ventral regions, separated by the
sulcus limitans, a longitudinal groove that forms as the neural tube matures. The dorsal
portion of the mature neural tube receives input from sensory neurons, while the ventral
portion is involved in motor functions. The spinal vertebrae are formed from the adjacent
somites, and the neural tube differentiates to the three tissue zones.
Cerebellar Organization. During cerebellar (cerebellum) organization, the three-

zone patterns are modified by cell migration, differential neuronal proliferation, and selective
cell death. A second mitotic layer, the external granular layer, forms at the region farthest
removed from the ventricular zone. Neuroblasts, or cells that will mature to neurons or glial,
from this layer will migrate back to the intermediate zone to form the internal granular layer.
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The original ventricular zone of the cerebellum creates a variety of neurons and glial cells,
including the Purkinje neurons, which are the major cell type in the cerebellum.
Cerebral Organization. Like the organization of the cerebellum, the three-zone

pattern is also modified during the organization of the cerebrum. The cerebrum is also
organized vertically into layers and the migrating neuroblast and glioblast form a cortical plate
containing six layers.
Figure 37. Differentiation of the Walls of the Neural Tube (Gilbert & Baressi, 2016)
Epidermis
The epidermis is derived from the ectodermal cells that surrounds and covers the embryo
after neurulation. This ectoderm is induced to form the epidermis than the neural tissues. During
development, the one-layered cell epidermis creates a second outer layer called the periderm.
The periderm covers the embryo; however, it sheds after the true epidermis has been formed
from the inner layer called the basal layer or stratum germinativum. The basal layer contains
the epidermal stem cell that are attached to the basement membrane. The epidermal stem cells
divide asymmetrically and the daughter cell that remains attached to the basal lamina remains a
stem cell, while the cell that migrates from the basal layer becomes differentiated to keratinocytes,
the primary cells found in the skin. In addition to the epidermis, cutaneous appendages such as
hairs, scaled and feathers are formed through inductive interactions between the dermal
mesenchyme and the ectodermal epithelium.
The Neural Crest
Sometimes called the fourth germ layer, the neural crest gives rise to several derivatives,
due to the population of multipotent progenitor cells that can produce diverse tissues. The table
below shows the different tissues or systems that originate from the neural crest.
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Some Derivatives of the Neural Crest (Gilbert & Baressi, 2016)

Peripheral Nervous System Neurons, including sensory ganglia, sympathetic and
parasympathetic ganglia, and plexus; neuroglial cells;
Schwann cells
Endocrine and Adrenal medulla; calcitonin-secreting cells; carotid body type I
Paraendocrine Derivatives cells
Pigment Cells Epidermal pigment cells
Facial Cartilage and Bone Facial and anterior ventral skull cartilage and bones
Connective Tissue Corneal endothelium and stroma; tooth papillae; dermis,
smooth muscle, and adipose tissue of skin, head, and neck;
connective tissue of salivary lachrymal, thymus, thyroid, and
pituitary glands; connective tissue and smooth muscle in
arteries of aorctic arch origin
Paraxial and Intermediate Mesoderm
In this section, the development of the mesodermal and endodermal germ layers will be
discussed. Generally, the mesoderm forms all the organs between the ectodermal wall and the
endodermal tissues. A neurula has a trunk mesoderm that can be divided into five regions: (1)
chordamesoderm, or the central region of trunk mesoderm, which forms the notochord, (2) the
paraxial, or somitic, mesoderm, which will form the somite, (3) the intermediate mesoderm,
which will form the urogenital system, (4) the lateral plate mesoderm, which gives rise to the
circulatory system, linings of the body cavities, and all mesodermal components of the limbs
except the muscles, and (5) the head mesoderm, which will form the connective tissue and
musculature of the head.
Figure 38. Major Lineages of the Amniote Mesoderm (Gilbert & Baressi, 2016)
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Paraxial Mesoderm
The somite derived from the paraxial mesoderm have cells that are committed only after
the somite is formed. As the somite matures, portions of it undergo epithelial-to-mesenchymal
transition, with cells that have restricted to form to only certain cell types, creating two major
compartments: the sclerotome and the dermomyotome. The table below shows the derivatives of
the somite.
Derivatives of the Somite (Gilbert & Baressi, 2016)

Traditional View Current View
Dermomyotome
Myotome forms skeletal Lateral edges generate primary myotome that forms muscles
muscles
Dermatome forms back Central region forms muscle, muscle stem cells, dermis, brown
dermis fat cells.
Sclerotome Forms vertebral and rib cartilage
Forms vertebral and rib Dorsal region forms tendons (syndetome)
cartilage Medial region forms blood vessels and meninges
Central mesenchymal region forms joints (arthrotome)
Forms smooth muscle cells of dorsal aorta
Intermediate and Lateral Plate Mesoderm
The intermediate mesoderm is responsible in forming the urogenital system, which

include the kidneys, the gonads and the associated cuts, and the adrenal gland. On the other
hand, the lateral plate mesoderm located at the farthest part away from the notochord forms the
circulatory system, including the heart, the blood vessels, and the blood cells. In addition, the
lateral plate mesoderm also lines the body cavities and the pelvic and limb skeleton.
Formation of the Kidney. The development of the mammalian kidney occurs within
three stages: (1) the development of the pronephric duct (day 22 in humans; day 8 in mice)
formed in the intermediate mesoderm, and the formation of the pronephros, or tubules of the
initial kidney, induced by the anterior region of the pronephric duct; (2) degeneration of pronephric
tubules and the induction of the remaining duct, nephric, or Wolffian duct, to form a new set of
tubules called the mesonephros, which is essential form blood cell development and tubules of
the male reproductive system (in mammals); and (3) the formation of the metanephros,
permanent kidney in amniotes, from interactions between epithelial and mesenchymal
components of the intermediate mesoderm.
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Figure 39. General Scheme of Development in the Vertebrate Kidney (Gilbert & Baressi,
2016)
Formation of Body Cavities and Linings. Body cavities and linings come from the
lateral plate mesoderm located at the lateral side of each of the two bands of intermediate
mesoderm. These lateral plates are split horizontally into two layers, with the dorsal layer,
somatic (parietal) mesoderm, and the underlying ectoderm forming the somatopleure. The
ventral layer, on the other hand, is called the splanchnic (visceral) mesoderm, which overlies
the endoderm and together with the endoderm, forms the splanchnopleure. Coelom, or the body
cavity, is formed in the space between the two layers. Later in development, the coelom divides
into two separate cavities, however, in mammals, mesodermal folds divide the coelom into the
pleural, pericardial, and peritoneal cavities, which will envelope the thorax, heart and abdomen,
respectively.
Formation of the Heart. The circulatory system is the first functional unit in the
developing embryo, responsible in providing nourishment, and the heart is the first functional
organ. The vertebrate heart arises from two regions of the splanchnic mesoderm. In mice, the
progenitor cells of the heart are already present during early amniote gastrula, and they are
located at the epiblast. These cells migrate together through the primitive streak and forms two
groups of lateral plate mesoderm cells. General specification of the cardiogenic mesoderm, or the
heart field, is already determined during this cellular migration. The figure below shows the
formation of the heart from heart fields in the mouse embryo.
Figure 40. The Heart Fields in a Mouse Embryo (Gilbert & Baressi, 2016)
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On the embryonic day 7.5, the heart fields from each side of the body have joined
into a common cardiac crest that contains the first and second heart fields. The first heart
field contributes primarily to the left ventricle. By day 10.5, the second heart field contributes
to the other three chambers—the right ventricle and the left and right atria—as well as to the
outflow tract that originally includes the aorta and the pulmonary artery.
Formation of the Blood Vessel and Blood cells. Along with the development heart,
the first circulatory loops are also formed. The development of blood vessels include two
processes: vasculogenesis and angiogenesis. Vasculogenesis creates networks of blood
vessels from the lateral plate mesoderm. Vasculogenesis occurs at different sites in
amniotes: (1) extraembryonic vasculogenesis, which occurs in the blood islands of the
yolk sac, and (2) intraembryonic vasculogenesis, which forms the dorsal aorta. On the
other hand, angiogenesis remodels this network and prune it into a distinct capillary bed,
arteries and veins. Hematopoiesis is the process of blood cell generation, and involves the
pluripotent hematopoietic stem cell, or hematopoietic stem cell, which can differentiate
to all blood cells and lymphocytes of the body.
Endoderm
The endoderm has two function in an embryo: (1) to induce the formation of several
mesodermal organs, and (2) construct the linings of the two tubes within the vertebrate body,
which are the digestive tube and the respiratory tube. In forming the anterior endoderm
portion of the digestive and respiratory tube, the embryo initially creates four pairs of
pharyngeal pouches with four pharyngeal arches in between. These structures will give rise
to several tissues: the first pair of pharyngeal pouches will become the auditory cavities of
the middle ear and eustachian tubes; the second pair of pouches gives rise to the walls of
the tonsils; the third pair of pouches becomes the thymus, and the one pair of the parathyroid
glands with the other pair derived from the fourth pair of pouches. The thyroid gland will be
formed from the midline of the pharynx. The figure below shows the formation of these
structures from the pharyngeal pouches.
Figure 41. Formation of the Glandular Primordia from the Pharyngeal Pouches
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The digestive tube located at the posterior to the pharynx constricts to form the future
pathway for food digestion – esophagus, stomach, small intestine, and large intestine. The
muscle used in peristalsis is derived from the mesenchyme cells from the splanchnic portion
of the lateral plate mesoderm. The linings of the liver, pancreas, and gall bladder are also
formed by the endoderm. On the other hand, the lungs are derived from the digestive tube.
References
Associates Inc.
Self Help.
Hall, B. K. (2015). Bones and cartilage: Developmental and evolutionary skeletal biology.
Stoddart, M. J., Craft, A. M., Pattappa, G., & Gardner, O. F. W. (Eds.). (2018). Developmental
biology and musculoskeletal tissue engineering : Principles and applications.
Let’s Check!
Activity 8. Now, that you know the basics of the processes of later embryonic development,
let us try to check your understanding of these topics. Match the structure and the germ layer
they are derived. Write your answers in the space provided:
_____ 1. Reproductive System a. Ectoderm
_____ 2. Urinary System b. Mesoderm

_____ 3. Digestive System c. Endoderm
_____ 4. Circulatory System
_____ 5. Brain
_____ 6. Spinal Cord
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_____ 7. Thyroid gland

_____ 8. Spleen
_____ 9. Epidermis
_____ 10. Vertebrate Eye
Let’s Analyze!
Activity 8. Now that you are acquainted with the processes during later embryonic
development, it is important that you will be able to relate this in your future profession. I will
1. Give examples of disorders that results from abnormalities in ectodermal
development.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Give examples of disorders that results from abnormalities in mesodermal

development.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. Give examples of disorders that results from abnormalities in endodermal

development.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 8. Based on your learnings about processes of later embryonic development, feel
free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
Keyword Index
Organogenesis Commitment
Competency Potency
Specification Stem cell
Differentiation Pre-cursor cells
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Big Picture
DEVELOPMENT OF THE TETRAPOD LIMB AND SEX DETERMINATION

a. Discuss the development of the tetrapod limb.
b. Discuss both chromosomal and environmental sex determination, and mammalian
gametogenesis

ULO-a. Discuss the development of the tetrapod limb.
Metalanguage

1. Tetrapods are animals with four limbs.

2. Stylopod is the most proximal region of the tetrapod limb and is adjacent to the
body wall.
3. Zeugopod is found in the middle region of the tetrapod limb.
4. Autopod is the most distal region of the tetrapod limb.
5. Pattern formation is a set of processes in which embryonic cells form spatial
arrangement of differentiated tissues.
6. Limb field is a region that have the cells capable of forming the limb.
7. Limb bud is a circular bulge containing mesenchymal cells capable of forming
the limb.
8. Apoptosis is the programmed death in cells.
9. Chondrocytes are cells found in the cartilage. They maintain and produce the
cartilaginous matrix.
10. Articulate cartilage is a kind of cartilage found in joints. It is a tough, elastic
fibrous connective tissue.
11. Synovium secretes the synovial fluid in joints to create lubrication.
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Key Knowledge
Animals that have four functional limbs are called tetrapod. The tetrapod limbs have
several functions. In most animals, limbs play a significant role in supporting the body,
locomotion, grasping, and many more. The bones of any tetrapod limb consist of a proximal
region called the stylopod, middle region, the zeugopod, and distal region called the
autopod. In animals, stylopod may be the humerus or femur, zeugopod may be the radius-
ulna or tibia-fibula, and the autopod may be the carpals-fingers or tarsal-toes.
There are three-dimensional coordinate systems in which a vertebrate limb functions:

(1) proximal-distal axis (shoulder-finger, or hip-toe), (2) anterior-posterior axis (thumb-
pinkie) and (3) dorsal-ventral axis (knuckles-palms)
Formation of Limb Buds
Limb buds are formed from proliferating mesenchyme cells from the somatic layer
of the limb field lateral plate mesoderm and from the somites at the same level. This
accumulation of the mesenchymal cell under the ectodermal tissue is the limb bud. In land
vertebrates, four limb buds appear in each embryo. This process is majorly influenced by the
expression of Hox gene in the anterior-posterior axis. In addition, the Hox genes also play a
vital role in the specification of forelimb and hindlimb. Early limb bud formation also requires
the induction of Wnt proteins and fibroblast growth factors.
Proximal-Distal Axis of the Limb
The apical ectodermal ridge, or AER, is the major signaling center for the
developing limb. It arises from the induction of mesenchyme cells to the overlying ectoderm
as they enter the limb field. AER has three major function: (1) enables linear growth of the
limb proximal-distal (shoulder-finger), (2) ensures the generation of the anterior-posterior
(thumb-pinkie) axis, and (3) specify the dorsal-ventral (knuckle-palm), as well as the anterior-
posterior axes. Generally, AER ensures the sustainability of the outgrowth and development
of the limb. The extension along the proximal-distal part of the limb is made possible by the
progress zone (PZ) mesenchyme, also called undifferentiated zone. The FGF proteins are
essential in generating the proximal-distal axis of the vertebral limb.
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Figure 42. Early Chick Forelimb Bud, Showing Apical Ectodermal Ridge (Gilbert &
Baressi, 2016)
Anterior-Posterior Axis of the Limb
The specification of the anterior-posterior axis of the limb is determined by a small

block of mesodermal tissues close to the posterior junction of the young limb bud and the
body wall. A region in the mesenchyme called the zone of polarizing activity (ZPA) induce
signals that instruct the limb bud to form along the anterior-posterior axis. The ZPA is defined
by its production of the sonic hedgehog protein. The sonic hedgehog protein specifies digit
identities. The cells producing the protein do not undergo apoptosis, or programmed cell
death, like the cells in the AER when they are finished with their job but becomes the bone
and the muscle of the posterior limb. In mouse, the digit 4 and 5 of its hindlimb arise from the
sonic hedgehog-producing cells.
Dorsal-Ventral Axis of the Limb
Determining the dorsal-ventral axis of the limb, that is knuckles and nails-pads and
soles, involves the ectoderm enclosing the tissue. The major molecule involved in the
generation of the dorsal-ventral axis of the limb is the Wnt7a, produced by the Wnt7a gene.
In chicken and mouse limb buds, this molecule is expressed in the dorsal ectoderm, and not
in the ventral side. Removing this molecule will result to ventral footpads on the surfaces of
the subject’s paws.
Formation of Digits and Joints in the Autopod
In sculpting the tetrapod limb, cell death, or apoptosis, plays an essential role of
separating our fingers and forming the joints. There are four major zones in which cells
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undergo apoptosis during the formation of the digits and the joints: (1) interdigital necrotic
zone, (2) interior necrotic zone, which separates the ulna and radius, (3 and4) anterior
and posterior necrotic zones, which shapes the end of the limb. Apoptosis in these zones
within the autopod is influenced by BMP proteins. BMP proteins are actively suppressed by
Noggin protein created in developing cartilage of the digits.
Figure 43. Patterns of Cell Death in leg primordia in duck and chicken (Glibert and
Baressi, 2016)
The formation of other joints in the body is also influenced by the BMP proteins. Aside
from inducing apoptosis, BMP proteins also play an important part in the formation of
cartilage and bone tissue by the induction of mesenchymal cells to become cartilage-
producing chondrocytes. The BMP proteins, BMP2 and GDF5 are expressed in regions
between the bones where the joints will form. The formation of the cartilage is further
promoted by the BMP7 protein. Wnt proteins also play a role in forming joints by sustaining
the expression of the said proteins during the formation. It is also important to note that
cartilage formation will not begin when there is the presence of blood vessels. Cells involved
in joint formation lose their chondrocyte characteristics to become articulate cartilage and
the synovium.
References
Associates Inc.
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Self Help.
Hall, B. K. (2015). Bones and cartilage: Developmental and evolutionary skeletal biology.
Let’s Check!
Activity 9. Now, that you know the basics of the development of the tetrapod limb, let us try
to check your understanding of these topics. Match the proteins/parts used according to the
corresponding axes established. Write your answers in the space provided:
_____ 1. Knuckles-palm a. Proximal-Distal
_____ 2. BMP proteins b. Anterior-Posterior
_____ 3. Wnt proteins c. Dorsal-ventral
_____ 4. Shoulder-finger
_____ 5. Hip-toe
_____ 6. Sonic Hedgehog protein
_____ 7. Thumb-pinkie
_____ 8. GDF5 proteins
_____ 9. FGF proteins
_____ 10. Hox gene
Let’s Analyze!
Activity 9. Now that you are acquainted with the basics of the development of the tetrapod
limb, it is important that you will be able to relate this in your future profession. I will require
you to give what are asked:
1. What is the difference of a duck’s webbed feet from a chicken’s feet in terms of
development?
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__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. What is the cause of polyploidy in humans? Explain its development.

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. What is the role of growth hormone and estrogen receptors in the later development
of humans?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What is the cause of dwarfism in humans? Explain its development.

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. What is the cause of gigantism in humans? Explain its development.

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 9. Based on your learnings about the basics of the development of the tetrapod limb,
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
Keyword Index
Tetrapod Autopod
Zeugopod Limb bud
Stylopod Necrotic zones
Axis Apoptosis
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ULO-b. Discuss both chromosomal and environmental sex determination, and
mammalian gametogenesis.
Metalanguage

1. Sex is the classification of male and female assigned during birth.

2. Bipotential, or indifferent, gonad is the common precursor for both ovaries and
testes.
Key Knowledge
In nature, there are two ways to determine the sex of an organism: (1) by
chromosomes and (2) influence of the environment. Some animals with sex determination
by chromosomes are mammals, birds, and Drosophila. On the other hand, sex in reptiles,
worms, and fishes, are often determined by different factors present in their environment.
Chromosomal Determination of Sex
Sex in mammals is often determined by the presence of either the second X, or Y

chromosomes in the 23rd chromosomal pair (female is XX; male is XY). In birds, female has
two dissimilar sex chromosomes (ZW), while male has similar sex chromosomes (ZZ). Flies
have the number of X chromosomes to determine the sex of the individual.
Mammalian Sex Determination
Mammals undergo two sex determination within their life cycle: primary and
secondary sex determination. During primary sex determination, the types of gonad
(ovaries or testes) are determined according to the sex chromosome received from the sperm
during fertilization. Both male and female gonads separate from a common precursor called
the bipotential, or indifferent, gonad. It is necessary to note that each haploid egg cell
contains a single X sex chromosome, while a sperm can carry either a single X, or Y, sex
chromosome. In addition, The Y chromosome is the determining factor in mammals, as it
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contains the genes that codes for a testis-determining factor, responsible in organizing the
gonad into a testis rather an ovary. The figure below shows the development of the testis
and the ovaries from an indifferent gonad.
Figure 44. Differentiation of the human gonads in transverse sections (Gilbert &
Baressi, 2016).
The proteins involved in the differentiation of the human gonads are the Wnt4, Sry
gene, Wt1, Lhx9, GATA4, and Sf1. The figure below shows the mechanism of primary sex
differentiation in mammals with respect to the expression of the mention proteins.
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Figure 45. Possible Mechanism for the Initiation of Primary Sex Determination in
Mammals (Gilbert & Baressi, 2016)
Secondary sex determination involves the determination of the sexual phenotype

outside the gonads, which include the internal and external reproductive system of both male
and female. This occurs during organogenesis of the embryo. Furthermore, another phase
of secondary sex determination also occurs later in the individual’s life, specifically during
puberty. The hormones released from the gonads determine the formation of the sexual
phenotype. The figure below shows the model for the formation of the external genitalia in
humans.
Figure 46. Model for the Formation of the External Genitalia in Humans (Gilbert &
Baressi, 2016)
In the model, the mesenchyme in the urogenital swelling secretes inhibitors of Wnt
signaling. Without the Wnt signaling, the estrogen modifies the genital tubercle into the clitoris
and the labioscrotal folds into the labia majora. On the other hand, the androgen in males
bind to the androgen receptor in the mesenchymal cells to prevent the synthesis of the Wnt
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inhibitors. The Wnt signaling is allowed, causing the formation of the penis and the
labioscrotal folds to become the scrotum from the genital tubercle.
Environmental Sex Determination
Factors in the environment that determines the sex in many organisms include
temperature, location, and the presence of other species member.
Temperature-Dependent Sex Determination
In reptiles, such as in crocodiles, sex determination is brought by the temperature of

eggs during a certain type of development. When incubated in low temperatures, these eggs
produce only either of the one sex. In contrast, incubation in high temperature results to the
opposite. The figure below shows the temperature-dependent sex determination in three
reptilian species. In the red-eared slider turtle (Trachemys scripta elegans), eggs incubated
at a temperature lower than 28°C will be males, while eggs incubate at a temperature higher
than 31°C produces female offspring.
Figure 47. Temperature-dependent Sex Determination in Three Reptilian Species

Location-Dependent Sex Determination
Some organisms that determine their sex according to the location are the echiuroid
worm Bonellia viridis and the snipper snail, Crepidula fornicata. In B. viridis, the larva
becomes a female if it lands in the ocean floor, however, if a larva is attracted to a female
proboscis, it enters the female’s body and becomes a sperm-producing symbiont in the
female. In C. fornicata, the position of a young snail in a mound made of snails determines
the sex of such snail. If the young snail is attached to a female, it becomes a male, however,
if the snail is removed from the mound then it becomes a female. Lastly, in fish populations,
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sex is often determined by the temperature and the number of male individuals present in
the population. A female can undergo sexual transition in the absence of a male through the
stimulation and increase of brain testosterone levels, which in turn, decrease the amount of
brain aromatase that often induce male development in other species of fish.
Mammalian Gametogenesis
Gametogenesis is the formation of gametes, the sperm, and the egg. The conserved
genes involved in the suppression of gene expression in germ cells include the (1) Vasa,
proteins involved in binding RNA and activation in germ-cell-specific messages; (2) Nanos,
proteins that bind to their partner, Pumilio, to form a very potent repressive dimer; (3) Tudor
proteins which affect sterility; and (4) Piwi, proteins involved in transcriptionally silencing
portions of the genome. In mammals, the cells where gametes are generated, the primordial
germ cells (PGCs), form in the posterior portion of the embryo before migrating into the
gonads. The newly formed PGCs enter the hindgut and are assisted by cells secreting stem
cell factor (SCF). The SCF is essential for PGC motility and survival. The gonad decides
whether the PGC becomes either sperm (spermatogenesis) or egg (oogenesis).
Sexual Dimorphism in Mammalian Meiosis (Gilbert & Baressi, 2016)
Meiosis, the process that reduces the number of chromosomes in germ cells to a
haploid complement in animals, occurs within the gonads. Meiosis differs from mitosis in that
meiotic cells undergo two cell divisions without an intervening period of DNA replication, and
the homologues chromosomes pair together and recombine genetic material. The first
meiotic division starts with a lengthy prophase, further subdivided into four stages. In
leptotene DNA replication has already occurred, and each chromosome consists of two
parallel chromatids. In this stage, the chromatin is stretched out very thinly, making it
impossible to identify each chromosome. The second stage, zygotene, is where synapsis
occurs. Synapsis is the pairing of homologous chromosomes together through the
synaptonemal complex. Tetrad or a bivalent refers to the complex of four chromatids and the
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synaptonemal complex. The third stage, the pachytene, is where crossing-over, or the
exchange of homologous genes between chromatids, occurs. Crossing-over may extend to
the last stage, the diplotene, but later, the synaptonemal complex breaks and homologous
chromosomes move away from each other but remain attached at various points called
chiasmata. During diplotene, there are high levels of gene transcription.
Meiotic metaphase (I) starts with diakinesis of the chromosomes. The chromosomes
migrate to form the metaphase plate after the nuclear envelope breaks down. In anaphase
I, the homologous chromosomes separate from each other, leading to telophase I, where
two daughter cells are formed, each with one partner of each homologous chromosome pair.
Second meiotic division occurs after a brief interkinesis. During meiosis II, the kinetochore of
each chromosome divides during anaphase resulting to new cells getting one of the two
chromatids. The result are four haploid cells that are genetically different.
Figure 45. Meiosis. (A) Prophase; and (B) Summarized First and Second Meiotic
Division (Gilbert & Baressi, 2016)
Spermatogenesis
Spermatogenesis is the developmental pathway to produce mature sperm from

germ cells. This begins during puberty and occurs in the recesses between the Sertoli cells.
Generally, spermatogenesis is divided into three phases: (1) proliferative phase, where
spermatogonia, or sperm stem cells, increase in number; (2) meiotic phase, which include
the two divisions creating haploid states; and (3) post meiotic shaping phase,
spermiogenesis, where round cells eject most of their cytoplasm and become a streamline
sperm.
The proliferative phase starts when the PGCs arrive at the genital ride of the male
embryo. These PGCs will become incorporated to the sex cords that will form the
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seminiferous tubules and will be called gonocytes. These gonocytes will become
undifferentiated spermatogonia and will be directly attached to the Sertoli cells through
adhesion molecules to be nourished.
Figure 46. Sperm Maturation. (A) Cross-Section of the Seminiferous Tubule; and (B)
Portion of Seminiferous Tubule showing Sperm Maturation (Gilbert & Baressi, 2016)
The cell division of these spermatogonia will form a population of differentiating

spermatogonia called Type A spermatogonia which can create more than 1000 sperm per
second in adult human males. Type B spermatogonia are further differentiated Type A
spermatogonia that responds to stem cell factors (SCF) and with high levels of Stra8
transcription factor (activated during puberty). They are precursors of spermatocytes and are
the last cells of the line that undergo mitosis. They divide once to produce primary
spermatocytes that enter meiosis. Each primary spermatocyte undergoes first meiotic
division to create a pair of secondary spermatocytes, called spermatids, which completes the
second division of meiosis. During the entire differentiation and meiotic process, the cells
move farther away from the basal lamina of the seminiferous tubule and closer to the lumen
where the spermatids differentiated into spermatozoa. The progression from the
spermatogonia to mature spermatozoa takes 65 days.
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Figure 47. Formation of Syncytial Clones of Human Male Germ Cells (Gilbert &
Baressi, 2016)
After the meiotic phase, spermiogenesis prepares the mammalian haploid

spermatid for functions of motility and interaction. Some of the preparations include the
construction of the acrosomal process with the acrosomal vesicles and the remodeling of
nucleosomes before the sperm enters the lumen of the seminiferous tubules. When the
sperms reside in the epididymis from the seminiferous tubules, they continue to develop. The
epididymal cells secrete exosomes, containing small ncRNAs and other factors that activate
and repress gene expression, which fuses to the sperm. Final differentiation of the sperm
occurs in the female reproductive tract in a process called capacitation.
Oogenesis
Oogenesis refers to the production of eggs. Mammalian egg maturation can be

divided into four stages: (1) proliferation of PGCs (will be called oogonia) in the ovary of the
human embryo, (2) the formation of primary oocytes after the first meiotic division, (3) the
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dictyate resting stage, where the primary oocytes remain is in the diplotene stage of the
first meiotic prophase (lasts from 12 to 40 years), and (4) continuation of the meiotic
division of oocytes when puberty occurs. After the first meiotic phase of the oocytes, one
of the two daughter cells contains hardly any cytoplasm, while the other daughter cell retains
nearly all the volume of cellular constituents. The smaller cell becomes the first polar body
while the larger cell is referred as the secondary oocytes. In the second meiotic division,
most of the cytoplasm is retained by the mature egg called the ovum, while the second polar
body forms, receiving little more than a haploid nucleus. In humans, however, the first polar
body dies within the first 20 hours after the first meiotic division. Thus, conserving the volume
of the oocyte cytoplasm in a single cell, rather than splitting it equally among four daughter
cells is the priority of oogenic meiosis.
References
Associates Inc.
Self Help.
Sex determination in vertebrates. (2019). ProQuest Ebook

Let’s Check!
Activity 10. Now, that you know the basics sex determination based on chromosome and the
environment, let us try to check your understanding of these topics. Match the following
organisms below to their mechanism of sex determination.
__________1. Fishes a. Chromosomal

__________2. Mammals b. Environmental
__________3. Insects
___________4. Reptiles
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___________5. Amphibians
___________6. Avian
Let’s Analyze!
Activity 10. Now that you are acquainted with sex determination based on chromosome and
the environment, it is important that you will be able to relate this in your future profession. I
will require you to give what are asked:
1. How is sex determined in Drosophila species?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. What is androgen insensitivity syndrome?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. What is pseudohermaphroditism?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What is the function of estrogen in mammalian sex determination?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. Give other disorders of sex chromosomes in humans.

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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In a Nutshell…
Activity 10. Based on your learnings about sex determination based on chromosome and the
environment, feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
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5.
Keyword Index
Temperature-dependent Sex
Sex determination
Determination
Primary Sex Determination Meiosis
Secondary Sex Determination Spermatogenesis
Location-dependent Sex Determination Oogenesis
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Big Picture
POSTEMBRYONIC DEVELOPMENT

a. Discuss the process of metamorphosis and its significance in some animals.
b. Discuss regeneration and its significance.
c. Discuss aging and senescence and its significance.

ULO-a. Discuss the process of metamorphosis in some animals.
Metalanguage

1. Metamorphosis is the hormonal reactivation of development that results to

giving an organism a new form.
2. Urodeles are amphibians belonging to the order Urodela. They are characterized
by their lizard-like appearance and having tail all their lives. Examples are newt
and salamanders.
3. Anurans are amphibians belonging to the order Anura and are characterized as
tailless. Examples of anurans are toad and salamanders.
4. Ammoniotelic animals excrete ammonia. Examples are fishes and tadpoles.
5. Ureotelic animals excrete urea instead of ammonia.
6. Molt, also known as sloughing and shedding, is the manner of which an animal
casts off a part of its body. This part may be, but not always, an outer layer or
covering. It may occur at specific times of the year, or a certain point in their life
cycle.
Key Knowledge
Development do not stop after embryonic development occurs. Organisms continue

to develop as they hatch or born and become mature enough to be able to reproduce and
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produce a new generation that will continue their existence. Members of the Animal Kingdom
can be divided into two types of developers: (1) direct developers, and (2) indirect
developers. Direct developers are animals that have their embryos develop into mature
individuals without involving a larval stage. Examples of these organisms are fishes, reptiles,
birds and mammals. On the other hand, indirect developers are animals that include sexually
immature larval stages that undergo metamorphosis, or hormonal reactivation of
development which gives the organism a new form. Typically, the larva has different food
requirements as compared to the adult form.
Amphibian Metamorphosis
Amphibians have a known life cycle that includes both water and terrestrial
environment (Greek amphi means double, and bios means life). For instance, tadpole, a
frog’s larva, sustains it growth and development in an aquatic environment and it undergoes
metamorphosis to become a fully mature adult frog capable of inhabiting the land. A frog
tadpole can be considered as a secondary larvae because its body plan is still the same
with an adult frog. On the other hand, primary larvae have body plans that are different from
the adult, such as in sea urchins.
Amphibians have two known groups – the anurans (frogs and toads) and the
urodeles (salamanders) – and both groups’ metamorphological process is initiated by the
thyroid hormones, such as thyroxine (T4) and tri-odothyroxine (T3). These hormones are
released in the blood throughout the organism’s body to initiate growth, cell death,
remodeling and respecification.
Growth of new structures in amphibians is largely influenced by T3. Examples of

structures developed during growth are the limbs, eye, eyelids and nictating membrane, and
neurons that serve on these organs. Cell death during metamorphosis is also influenced by
T3 which causes the paddle-like tail and the gills of the tadpole (amphibian larva) to
degenerate. The larval gills degeneration is achieved due to the induction of the hormone to
the cells to commit apoptosis. On the other hand, the larval tail degenerates because of the
action of macrophages. Aside from these two structures, the tadpole’s red blood cells are
also replaced by adult red blood cells to accommodate the demands of living in a terrestrial
environment. Furthermore, some larval organs are remodeled during the course of
metamorphosis. For instance, the larval intestine, which is usually coiled because of a
herbivorous diet, is remodeled to become shorter to accommodate a carnivorous diet. The
larva’s nervous system and skull is also remodeled. Respecification includes biochemical
changes within amphibian’s body. In amphibians, the most dramatic change occurs in the
liver. Tadpoles are ammonotelic, in which they excrete ammonia. Adult frogs, such as the
genus Rana, excrete urea, and thus considered as ureotelic. During metamorphosis, the
liver starts to create enzymes to produce urea from carbon dioxide and ammonia. This
respecification is induced by T3. The table below show some metamorphic changes in
anurans.
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Some Metamorphic Changes in Anurans (Gilbert & Baressi, 2016)
Insect Metamorphosis
Insects have three kinds of development: (1) ametabolous development, where an

insect do not have a larval stage (ex. springtail), (2) hemimetabolous development, where an
insect undergoes a gradual or incomplete metamorphosis, mainly consisting of three life
stages: egg, nymph and adult (ex. cockroach), and (3) holometabolous development (ex.
butterfly and beetles), generally consisting of four life stages: egg, larva, pupa and adult.
Source: https://www.letstalkacademy.com/ckeditor/plugins/imageuploader/uploads/66231363a.jpg
Figure 48. Three Kinds of Development in Insects
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Ametabolous insects have a pronymph stage as the insect hatches from the
eggs. This pronymph resembles its adult counterpart and it grows with each molt with a
new cuticle. Hemimetabolous insects also have a brief pronymph stage followed by a
nymph stage, where the insects look like an immature adult. Rudiment structures, such
as the wings and genital organs and other adult structures, become present and will be
gradually developed into mature parts with each molt. At the final molt, a winged and
sexually mature adult, or imago, will emerge. Holometabolous insects have no
pronymph stage. The juvenile form that hatches from the egg is called a larva. Like the
other types of development, the larval also undergo molts enabling it to become larger.
The stages between these molts are called instar. Eventually, the larval will undergo a
metamorphic molt to become a pupa. Dramatic changes occur during this stage of
transition between the larva and adult stages. Imaginal molt then forms the adult
(imago) cuticle beneath the pupal cuticle. The adult emerges from the pupal case at adult
eclosion. Molting is caused by the hormone 20-hydroxyedcysone (20E). If there are
high levels of juvenile hormone, the molt gives rise to another larval instar. In low
concentrations, the molt produces a pupa. In the absence of the juvenile hormone, the
molt is an imaginal molt.
Source: https://lh3.googleusercontent.com/proxy/4nQTBu7MUYaUNveBmDca6DMa-B8KdYPvMJMCIUJfdo5Om7zS8dqTEjaLX4WS8Ixg1NTH6O1JAbmduNq4OxbXI6LlN6m7CjaUReUaCYuFY-
l3Azs9dKWI1hG0VsNfaFx-bMIh1Qg
Figure 49. The Life Cycle of a Coconut Rhinoceros Beetle
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References
Associates Inc.
Self Help.
Heming, B. S. (2018). Insect development and evolution. ProQuest Ebook

Echinoderms. (2019). ProQuest Ebook Central https://ebookcentral.proquest.com
Hoy, M. A. (2018). Insect molecular genetics : An introduction to principles and applications.

Let’s Check!
Activity 11. Now, that you know some basic concepts of metamorphosis in animals, let us try
to check your understanding of these topics. Determine the kind of developer for the animals
stated below:
__________1. Butterfly a. Indirect
__________2. Cockroach b. Direct

__________3. Humans
__________4. Snails
__________5. Tunicates
Determine the type of insect development for the following insects:

__________6. Cicada c. Ametabolous
__________7. Grasshopper d. Hemimetabolous
__________8. Dragonfly e. Holometabolous

__________9. Bees
__________10. Flea
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Let’s Analyze!
Activity 11. Now that you are acquainted with some basic concepts of metamorphosis in
animals, it is important that you will be able to relate this in your future profession. I will require
1. What is the evolutionary significance of metamorphosis?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Briefly discuss of the metamorphosis of the pluteus larva.

__________________________________________________________________
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__________________________________________________________________
3. What happens if the thyroid gland is removed in an amphibian tadpole?

__________________________________________________________________
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__________________________________________________________________
4. What is a threshold model in amphibian metamorphosis?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. What is the significance of remodeling the nervous system during metamorphosis in

amphibians?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 11. Based on your learnings about some basic concepts of metamorphosis in animals,
1. __________________________________________________________________
__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
Keyword Index
Metamorphosis Larva
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Holometabolous Ametabolous
Hemimetabolous Direct and Indirect Developers
Anurans Urodeles

ULO-b. Discuss regeneration and its significance.
Metalanguage

1. Regeneration is a postembryonic reactivation of development to restore missing

or damaged tissues.
2. Transdifferentiation is a process in which one type of adult tissue differentiates
to another type of adult tissue. Morphallaxis (morphallactic regeneration) is a
mode of regeneration by the transdifferentiation of existing body tissues.
3. Epimorphosis (epimorphic regeneration) is a mode of regeneration involving
extensive cell proliferation followed by differentiation. A blastema is a mass of
cells formed in epimorphosis capable of growth and regeneration.
4. Hydra are cnidarians belonging to the class Hydrozoa. They are invertebrates
characterized by their capability to completely regenerate from fragmentation.
5. Flatworms or platyhelminths is a phylum of invertebrates that are characterized
by their flat bodies. There are four known classes of flatworms: flukes
(Trematoda), tapeworms (Cestoda), Monogena, and planarians (Turbellaria).
Key Knowledge
Regeneration is the replacement of a body parts one the original parts have been
remove. Some groups in the Animal Kingdom possess this ability, having the capability to
restore parts amputated or grow an entire organism from small body fragment. In humans,
however, regeneration of an entire individual is not yet possible. The figure below shows
some representative organisms and their regenerative capabilities.
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Figure 50. Representative Organisms and Their Comparative Regenerative

Capabilities (Gilbert & Baressi, 2016)
There are four mode of regeneration: (1) Stem-cell mediated regeneration, (2)
epimorphosis, (3) morphallaxis, and (4) compensatory regeneration. Stem-cell
mediated regeneration is mode of regeneration that allows an organism to regrow
certain tissues or organs that have been lost from stem cells. Example of this is the
continuous replacement of blood cells from hematopoietic stem cells residing in the bone
marrow. Epimorphosis is a regeneration mode where adult structures undergo
dedifferentiation to form an undifferentiated cell mass called a blastema. The blastema
can undergo redifferentiation to form new structures. Epimorphosis is evident in
regenerating amphibian limbs. Morphallaxis involves the repatterning of existing tissues.
This can be referred to as transdifferentiation and can be seen in hydra. Lastly,
compensatory regeneration involves the division of differentiated cells while retaining
their differentiated state and functions. This is evident in mammalian liver. The figure
below shows the four models of regeneration.
Figure 51. Four modes of regeneration (Gilbert & Baressi, 2016)

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Hydra. Hydra is a genus of cnidarians. They are known for their capability of
stem-cell mediated regeneration, morphallaxis and epimorphosis. Hydras are
simple in their morphology – they have a tubular, radially symmetric body with a
head in its distal end a foot at its proximal end. The head consists of a conical
hypostome, containing the mouth and a ring of tentacles for eating and foraging,
while the foot, or also called the basal disc, is responsible for sticking to rocks and
undersides of pond plants. Naturally, hydra reproduces through sexual and asexual
reproduction.
Stem-cell mediated regeneration in hydra is evident during its routine cell

replacement by three types of stem cells: (1) unipotent endodermal cells, (2)
unipotent ectodermal cells, and (3) multipotent interstitial stem cell located at the
ectodermal layer. The interstitial stem cell is capable of generating neurons,
secretory cells, nematocytes and gametes. Both morphollaxis and epimorphosis
occur when some parts of the hydra is removed. When a hydra loses its head, the
protein family Wnt is activated at the apical portion to form the new head. However,
if the cut is just below the hypostome, a specific Wnt protein, Wnt3 protein, is
upregulated in the epithelial cells near the cut surface to induce remodeling of
existing cells to form the head. Because there is no proliferation of cells during this
process, it is called morphallactic regeneration (regeneration by cell
transdifferentiation). If a hydra is cut at its midsection, the cells derived from the
interstitial stem cell release Wnt3 proteins and undergo apoptosis. The burst
production Wnt3 triggers the beta-catenin protein in the interstitial cells beneath
them that causes the proliferation in the interstitial cells and remodeling in the
epithelial cells, thus, resulting to epimorphic regeneration or epimorphosis.
Flatworms. Planarian flatworms, or free-living flatworms, are capable of

stem-cell mediated regeneration achieved by pluripotent stem cells called
clonogenic neoblasts (cNeoblast). cNeoblast are set of pluripotent cells that serve
to replace aging cells of the flatworm adult body. In addition, these cells can also
migrate to wound sites and regenerate the tissue. When the flatworm is cut in half,
the anterior portion of the lower half generates a new head while the posterior of the
upper half regenerates a tail. If a flatworm is cut into three pieces, the middle piece
will regenerate a head from its anterior end and a tail in its posterior end. However,
if the middle slice is too narrow, regeneration is abnormal. The figure below shows
flatworm regeneration and its limits.
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Figure 52. Flatworm regeneration and its limits (Gilbert & Baressi, 2016)
Salamanders. Salamanders are capable of epimorphic limb regeneration,

that is, when limbs are amputated, the remaining limb cells reconstructs a new limb.
Through epimorphic regeneration, a regeneration blastema, an aggregation of
undifferentiated cells derived from the original differentiated tissues, proliferates and
redifferentiate to new limb parts. The figure below shows regeneration of
salamander forelimb.
Figure 53. Regeneration of a Salamander Forelimb (Gilbert & Baressi, 2016)
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Mammals. In mammals, such as in rodents and humans, the tips of the digits
can regenerate if the organism is young enough. This is possible because of
regeneration blastema composed of progenitor cells forming at the tips of the digits.
Furthermore, mice heart tissue can also regenerate, however, it is only restricted
during the first week of neonatal life. In humans, the liver can regenerate through
compensatory regeneration, or the division of differentiated cells to recover the
structure and function of the organ. This is also true with the zebrafish heart.
References
Associates Inc.
Self Help.
Epigenetics and regeneration. (2019). ProQuest Ebook

Mechanisms of regeneration. (2014). ProQuest Ebook

Let’s Check!
Activity 12. Now, that you know some basic concepts of regeneration in animals, let us try to
check your understanding of these topics. Determine what mode of regeneration is asked for
the following below:
__________1. Blood cell replacement a. stem-cell mediated regeneration
__________2. Skin cell replacement b. epimorphosis
__________3. Tail regeneration in salamander c. morphallaxis

__________4. Wound healing in humans d. compensatory regeneration
__________5. Regeneration of liver
__________6. Regeneration of hydra from a fragment
__________7. Regeneration of starfish from a fragmented limb

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___________8. Limb regeneration in crabs

___________9. Tapeworm body regeneration from head
__________10. Tail regeneration in lizards
Let’s Analyze!
Activity 12. Now that you are acquainted with basic concepts of regeneration in animals, it is
important that you will be able to relate this in your future profession. I will require you to give
what are asked:
1. Why is it possible for a human to survive after giving a portion of his liver in a
transplant?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. What are other human body parts that can regenerate?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. What is autotomy and its significance? Which animals are capable of autotomy?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What is hyperplasia and its role in regeneration in humans?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. Tapeworms continue to thrive within a human body as long as its head is still present.
How is this possible?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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In a Nutshell…
Activity 12. Based on your learnings about the basic concepts of regeneration in animals,
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
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Keyword Index
Regeneration Blastema
Transdifferentaition Compensatory regeneration

Morphallaxis Stem-cell mediated regeneration
Epimorphosis cNeoblast

ULO-c. Discuss aging and senescence and its significance.
Metalanguage

demonstrate ULO-c will be operationally defined to establish a common frame of reference
1. Aging is the time-related decline and the deterioration of physiological functions

necessary for survival and fertility.
2. Senescence is the facet of aging that involves the gradual deterioration of
functional characteristics.
3. Maximum lifespan is another facet of aging that shows the maximum number of
years an individual of a given species has been known to survive and is a
characteristic of that species.
4. Life expectancy is a characteristic of a population that shows the average length
of time a given individual of a given species can expect to live.
5. Reactive oxygen species are unstable molecules, containing oxygen that easily
reacts with other molecules in a cell. The build-up of these molecules can cause
damage to proteins and nucleic acid and may even cause cell death
Key Knowledge
Genes Involved in Aging
The nature of genes among organisms determines the species’ longevity.

Across animal species, there are four sets of genes involved in the aging and its
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prevention. These genes are conserved between phyla and even kingdoms, which
means that it has been maintained in natural selection and cannot be altered by
evolutionary processes. Furthermore, these conserved genes are unique and essential.
The genes involved are (1) DNA repair enzymes, (2) proteins of the insulin signaling
pathway, (3) proteins in the mTORC1 signaling pathway, and (4) chromatin
remodeling enzymes.
DNA Repair Enzymes
DNA repair enzymes are enzymes able to recognize and correct physical
damages in DNA due to radiation, UV light or reactive oxygen species. Species with
efficient DNA repair enzymes are observed to have longer lives. The ‘wear-and-tear’
theories of aging proposed that small trauma to the human body and in its genome build
up as we age. This also mean that point mutations also increase as age increases and
the efficiency of the enzymes encoded by genes also decreases. Moreover, if mutations
occur in the genes which encode transcriptional or translational proteins, faulty proteins
within the cells will increase. Within the human body, the source of mutations includes
radiation and reactive oxygen species (ROS) produced by normal metabolism. ROS
such as superoxide ions, hydroxyl (‘free’) radicals, and hydrogen peroxide, can oxidize
and do significant damage in cell membranes, proteins, and nucleic acid. Ultraviolet
radiation, oxidative stress and other factors that can damage the DNA, also cause a
transcription factor, p53, to separate from its repressor, allowing it to function. p53 can
halt cell cycle, cause cellular senescence, initiate cellular apoptosis, and activate DNA
repair enzymes, which means that its activation can either be beneficial or deleterious.
Other method of activating the p53 is by destroying the telomeres, the protective
nucleoprotein caps on chromosomal tips. There is a positive correlation between
telomere length and longevity in humans. The enzyme telomerase maintains telomere
integrity, acting as an antisenescence complex.
Source: https://microbewiki.kenyon.edu/images/thumb/d/dc/Img-research-Telomerase_Genomics-illiustration.png/300px-Img-research-Telomerase_Genomics-illiustration.png
Figure 54. Telomeres and Telomerase in Chromosomes

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Aging and Insulin Signaling Cascade
In some species, such as in Caenorhabditis elegans, the organism can extend

its lifespan by becoming metabolically dormant during the larval stage. This stage is
called the Dauer larva stage, a nonfeeding state of diapause, a condition in which
development and aging are suspended. This is made possible by the regulation of the
insulin signaling pathway in nematodes (DAF-2 pathway), and only occurs if the
nematodes are overcrowded or there is insufficiency of food. Downregulation of insulin
signaling pathway has several other functions: (1) influences metabolism, (2) increases
the cellular production of DNA repair enzymes and enzymes that prevent oxidative
damage, and (3) decreases fertility. In mammals, there are few evidence that the insulin
signaling pathway affect life span. In dogs, however, breeds with low levels of insulin-
growth factor 1 (IGF 1) live longer than breeds with higher levels of this factor. In
Drosophila, flies with weak loss-of-function mutations of the insulin receptor genes or
genes in the insulin signaling pathway live nearly 85% longer than wild-type flies,
however, these mutants are sterile, and their metabolism seemed to be in diapause.
Calorie restriction is way of downregulating the insulin pathway. In studies involving
primates (including human), calorie restriction appears to slow down age-associated
decline of heartbeat variability and motor coordination.
The mTORC1 Pathway
The mTORC1 pathway has four major functions: cell growth and proliferation,
angiogenesis, bioenergetics and cell survival through DNA repair, and decreasing
autophagy, the removal and replacement of damaged organelles and senescent cells,
and apoptosis. There is an inverse relationship between the number of mTORC1 with
the amount of autophagy.
Chromatin modification
The sirtuin gene prevents aging throughout the eukaryotic kingdom. This gene
encodes histone deacetylation (chromatin-silencing) enzymes, prevents gene from
being expressed when unnecessary, and repair chromatin breaks. When DNA strands
break due to aging, sirtuin proteins are needed to fix these DNAs, resulting to the
proteins not attending to their usual function. This results to the other silenced genes
becoming active as the cell ages. In mammals, it was shown that infusing inhibitors of
histone deacetylase can increase the ability to store memories in aged mice.
Exceptions to the Aging Rule
Some organisms have unique aging characteristics. In turtles, their mortality rate
does not increase with age, and their reproductive rate does not decrease with age
either. In monarch butterflies, their longevity is often affected by external factors such
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as the season. The juvenile hormone in monarch butterflies, when expressed, promotes
fertility but shorter longevity. When suppressed, such as in butterflies that migrate to
wintering grounds, the butterflies lose their fertility and extend their longevity. In
cnidarians, on the other hand, the hydra seems to be immortal because of their capacity
to retain their stem cell populations. In hydrozoans Turriptosis dohrinii and Hydractina
carnea, the medusae of these species can revert to it polyp state even after being
sexually matured and is called reverse development.
References
Associates Inc.
Self Help.
Musi, N., & Hornsby, P. (Eds.). (2015). Handbook of the biology of aging. ProQuest Ebook
Let’s Check!
Activity 13. Now, that you know some basic concepts about aging and senescence, let us try
to check your understanding of these topics. Give what is asked below.
_________1. These are hydrozoan species that can revert their medusae form back to the
polyp form.
_________2. This is a gene that helps prevent aging across the Domain Eukarya.
_________3. It is the time-related decline and the deterioration of physiological functions
necessary for survival and fertility.
_________4. It is the facet of aging that involves the gradual deterioration of functional
characteristics.
_________5. These are unstable molecules, containing oxygen that easily reacts with other
molecules in a cell.
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Let’s Analyze!
Activity 13. Now, that you know some basic concepts about aging and senescence, it is
important that you will be able to relate this in your future profession. I will require you to give
what are asked:
1. What happens to stem cells when a human age?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Give examples of antioxidants. How do antioxidants stop humans from manifesting

signs of aging?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. What is Hutchinson-Gilford syndrome?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What are the factors that shorten telomeres? What happens when the telomere
becomes too short?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. Why do you think the genes for aging and senescence are conserved all throughout
the evolutionary history?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 13. Based on your learnings about some basic concepts of aging and senescence,
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1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
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Keywords Index
Aging Reactive oxygen species
Senescence DNA repair enzymes

Maximum lifespan Telomeres
Life expectancy Juvenile hormones
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Big Picture
DEVELOPMENT IN HEALTH AND DISEASE, ENVIRONMENT, AND EVOLUTION

a. Discuss development in health and disease, including birth defects, endocrine
disruptors, and cancer.
b. Discuss development and environment, including biotic, abiotic and symbiotic
regulation.
c. Discuss development and evolution, including developmental mechanisms of
evolutionary change.

ULO-a. Discuss development in health and disease, including birth defects,
endocrine disruptors, and cancer.
Metalanguage

1. Stochastic means random.

2. Congenital abnormalities are defects present at birth.
3. Syndrome is a group of symptoms which consistently occur together.
4. Aneuploidy is the presence of an abnormal number of chromosomes in the cell.
5. Pleiotropy happens when one gene influence two or more seemingly unrelated
phenotypic traits.
6. Teratogens are any agents that can disturb or affect the development of an
embryo or a fetus.
7. Mosaic pleiotropy occurs when a single locus directly affects two phenotypic
traits, and the effects are produced independently because of the gene being
critical in different parts of the body.
8. Relational pleiotropy occurs when the action of a single locus initiates a
cascade if events that impact multiple independent traits.
9. Genetic heterogeneity occurs when mutations in different genes can produce
the same phenotype.
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10. Phenotypic heterogeneity occurs when same mutation can produce different
phenotypes in different individuals.
Key Knowledge
In animals, the major pathways that may lead to abnormal development: (1)
genetic mechanisms, (2) environmental mechanisms, and (3) stochastic (random)
events.
Genetic Errors of Human Development
Syndromes are several abnormalities occurring together. These syndromes may

be caused chromosomal events (for instance, aneuploidy) where several genes are
added or deleted, or the production of several effects caused by a single or a pair of
genes (pleiotropy). Syndromes can either be mosaic pleiotropy or relational pleiotropy.
Mosaic pleiotropy occurs when a single locus directly affects two phenotypic
traits, and the effects are produced independently because of the gene being critical in
different parts of the body. An example of mosaic syndrome is Down Syndrome (Trisomy
21) caused by an extra copy of the chromosome 21 (an aneuploidy). The over
production of transcription factors and regulatory microRNAs from the extra
chromosomes causes misregulation of genes necessary for heart, muscle, and nerve
formation.
Source: https://obgynkey.com/wp-content/uploads/2016/06/B9781437711554001176_f117-001-9781437711554.jpg
Figure 55. Trisomy 21 or Down Syndrome

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Relational pleiotropy occurs when the action of a single locus initiates a

cascade if events that impact multiple independent traits. An example for this is the
disease phenylketonuria, which is caused by a defect in the liver enzyme phenylalanine
hydroxylase causing excess plasma levels of phenylalanine. Consequently, the
myelination of the axons in the brain is affected and brain and ultimately cause mental
retardation.
Genetic heterogeneity occurs when mutations in different genes can produce

the same phenotype. This means that similar phenotypic result will occur regardless on
which gene in the same signal transduction pathway is affected. An example for this is
tuberous sclerosis, which is caused by mutations in either chromosome 16 or
chromosome 9 and retinitis pigmentosa, which has autosomal dominant and recessive
X-linked recessive forms. Phenotypic heterogeneity occurs when same mutation can
produce different phenotypes in different individuals. An example can be seen in
heritable mutations in the FGFR3 gene, a gene involved in forming bones. It was
observed that in a family, the phenotypic result of the mutation in an individual varies
and can range from severe phocomelia or lack of limbs to mild abnormality of the thumb.
Teratogenesis
Teratogens are any agents that can disturb or affect the development of an
embryo or a fetus. This may lead to a congenital abnormality in a child and can even
abort the pregnancy. Examples of teratogens are chemicals and drugs. The figure below
shows some teratogens that causes disruptions in human development.
Source: https://www.researchgate.net/profile/Zohaib-Sultan-4/publication/287194568/figure/tbl2/AS:669620507783184@1536661378600/Drug-and-maternal-teratogens-and-possible-undesired-effects-Known-side-effects.png
Figure 56. Example of Teratogens
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The human fetus is very sensitive to teratogens during weeks 3 to 8, also known
as the embryonic period. In this period most of the organ systems are still forming.
However, the fetal period (the remaining time in the utero) can still be a window for
functional defects and minor anomalies caused by teratogens.
Figure 57. Embryonic and fetal period and susceptibility to teratogens (Gilber
and Gilbert, 2016)
Endocrine disruptors are exogenous chemicals that disrupt development by

interfering with the normal functions of hormones. Most of the effects of endocrine
disruptors are only evident microscopically, but with major physiological changes. These
chemical factors can mimic the effect of a natural hormone, act as antagonist and inhibit
the binding of a hormone to its receptor or block hormone synthesis, affect the synthesis,
transportation, or elimination of a hormone in the body, or can induce sensitivity to
hormones in an organism’s later life. The table below shows chemical known to be
endocrine disruptors.
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Source: https://d3i71xaburhd42.cloudfront.net/fb318d6eddf791b5d3129f99fa2baca2f1145f4a/2-Table1-1.png
Figure 58. Example of Endocrine Disruptors
Cancer as a Disease of Development
Carcinogenesis are aberrations of the processes that underlie differentiation and

morphogenesis, making it as a disease of development. Both congenital anomalies and
carcinogenesis are often caused by defects in the same pathways. There several facets
in viewing malignancy and metastasis in terms of development including (1) context-
dependent tumor formation, (2) defects in cell-cell communication as the initiator of
cancer, (3) cancer stem cells, and (4) epigenetic reprogramming of cancer cells.
Context-dependent tumors. Tumors often become malignant due to

environmental factors. In an experiment in mice, teratocarcinoma, or tumor of germ cells
or stem cells, which often kills adult mice, lose their malignancy when placed in the inner
cell mass of a mouse blastocyst. The cells continue to divide normally, and its cellular
progeny become part of embryonic organs. It is assumed that the stem cell environment
suppresses tumor formation by secretion of inhibitors in the paracrine pathways.
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Defects in cell-cell communication. Cancer cells can be caused by

miscommunication between cells, which can occur by defects in tissue architecture. In
an experiment in rats, it was observed that epithelial cell cancers do not directly stem
from epithelial cells, but rather from the mesenchymal stromal cells that surround and
sustain the epithelia. The mesenchymal stromal cells, which have been treated with
carcinogens and have sustained defects, adversely affected the capacity of the cells to
instruct epithelial cells resulting to altered tissue architecture and a loose control of cell
proliferation of the latter.
Defects in paracrine pathways. Tumors can result from disruptions of

paracrine signals in between cells. For instance, tumor cells can secrete paracrine
factors such as the Sonic hedgehog (Shh) which can act as an autocrine factor that
stimulate the tumor cells to grow. Blocking the pathway for this paracrine factor can
reverse medulloblastomas and leukemias. In addition, the Shh can also act on other
cells, such as the stromal cells mentioned above, to secrete factors such as the insulin-
like growth factor (UGF) that supports the tumor cells. By blocking the pathway, the
tumor was observed to regress.
The cancer stem cell hypothesis. Cancer has stem cells that continues to
proliferate and differentiate. In glioblastomas, prostate cancer, melanomas and myeloid
leukemias, cancer stem cell (CSC) gives rise to more cancer stem cells and to
populations of relatively slowly dividing differentiated cells. Like normal stem cells, CSC
can self-renew and generate non-stem cell populations of the tumor.
Developmental Therapies for Cancer
One mode of treatment for cancer is differentiation therapy, where malignant

cells are encouraged to differentiate into more mature forms by using pharmacological
agents. It is anchored on the concept that malignant tumor cells tend to assume a less
specialized, stem cell-like dedifferentiated state.
References
Associates Inc.
Self Help.
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Newton, R. W., Marder, L., & Puri, S. (2015). Down syndrome Current perspectives.
Let’s Check!
Activity 14. Now, that you know some basic concepts about development in health and
disease, let us try to check your understanding of these topics. Give what is asked below.
___________1. This is an endocrine disruptor that causes vaginal cancer and reproductive
tract abnormalities.
___________2. This is an endocrine disruptor that causes acceleration of puberty in rodents.
___________3. This is an endocrine disruptor that causes delated puberty and reduced sex
accessory gland size in male rodents.
___________4. This is an endocrine disruptor that causes delayed neurological
development in human fetus.
___________5. This is an endocrine disruptor that can cause decrease in testosterone levels.
___________6. This is a teratogen that can cause cleft lip and palate.
___________7. This is a teratogen that can cause brain damage and microcephaly.
___________8. This is a teratogen that can cause liver damage in the fetus.
___________9. This is a teratogen that can cause maternal toxicity and discoloration of tooth.
___________10. This is a teratogen that can cause placental abruption and cognitive delay.
Let’s Analyze!
Activity 14. Now, that you know some basic concepts about development in health and
disease, it is important that you will be able to relate this in your future profession. I will require
1. What is fetal alcohol spectrum disorder?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. What is the mechanism of the endocrine disruptor diethylstilbestrol (DES)?

__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
3. What is epi-alleles? What role do epi-allele play in transgenerational inheritance of
development disorders?
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__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. Cancer is a disease of development. Elaborate.

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. What is differentiation therapy? How is it applied in Oncology?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 14. Based on your learnings about some basic concepts of development in disease
and health, feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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1.
2.
3.
4.
5.
Keywords Index
Teratogens Pleiotropy
Syndrome Endocrine disruptors

Congenital abnormalities Cancer
Aneuploidy Tumor

ULO-b. Discuss development and environment, including biotic, abiotic
and symbiotic regulation.
Metalanguage

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1. Phenotypic plasticity is an organism’s ability to react to environmental inputs by

changing its form, state, movement, or rate of activity.
2. Developmental plasticity, on the other hand, occurs during embryonic or larval
stages in animals, or even plants.
3. Morphs, or ecomorphs are different phenotypes bought by environmental
conditions.
4. Kairomones are chemicals from the predators that can induce the developing prey
to change its phenotype to defend itself.
Key Knowledge
Role of Environment in Producing Phenotypes

There are several ways in the environment can affect development and influence
phenotypes. Phenotypic plasticity is an organism’s ability to react to environmental inputs
by changing its form, state, movement, or rate of activity. Developmental plasticity, on the
other hand, occurs during embryonic or larval stages in animals, or even plants. There are
two types of phenotypic plasticity: reaction norms and polyphenism. Reaction norm, also
called a norm of reaction, is a pattern of phenotypic expression of a single genotype across
a range of environments. The different phenotypes bought by environmental conditions are
called morphs, or ecomorphs. On the other hand, polyphenism describes discontinuous
phenotypes elicited by the environment.
Source: https://image.slidesharecdn.com/hgmlfwxiqo2ysurjjdaf-signature-7770ee07ff1c3b2eb683c51e945fccf15e8e48aa392406033e715d2c9ad74a51-poli-
150307101220-conversion-gate01/95/dna-methylation-and-diet-9-638.jpg?cb=1425799821
Figure 59. Reaction Norm vs. Polyphenism
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Diet-induced polyphenism. In some organisms, diet plays an important role in

determining the phenotype of a developing animal. In the caterpillar Nemoria arizonaria,
caterpillars that hatch on oak trees during spring has a form that blends with the young oak
flowers, while on the summer, the caterpillars that hatches resembles newly formed twigs
instead. This can act as a mimicry defense for the caterpillar against predators.
Figure 60. Diet-induced Polyphenism in Nemoria arizonaria (Gilbert & Baressi, 2016)
In ants, wasps and bee colonies, larvae that are adequately fed become queens.
The protein that induces the transformation of the larvae to a queen is called royalactin.
Figure 60. Nutrition-induced Polyphenism in the ant Pheidologeton (left) and Apis
mellifera (right) (Gilbert & Baressi, 2016)
In male dung beetles (Onthopagus sp), the anatomical and behavioral phenotypes
are determined by the quality and quantity of the dung provided by the mother. The amount
and quality of food determines the titer of juvenile hormone during the larva’s molt, which in
turn, determine the larva’s size during metamorphosis and the growth of the imaginal disc
that makes the horn.
Predator-induced polyphenisms. Predator-induced polyphenism describes the
ability of an organism to modulate development in the presence of predators. Some predators
release chemicals called kairomones that can induce the developing prey to change its
phenotype. The figure below shows some examples of predator-induced polyphenism.
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Figure 61. Predator-induced defenses (Gilbert & Baressi, 2016)
Temperature as Environmental Agent. As discussed, temperature can determine

the sex in some species of organisms such as in cold-blooded vertebrates like fish, turtles,
and alligators. One of the major disadvantages of this temperature-dependent sex
determination is the narrowing of the temperature limits in which a species can exist. Due to
changing global temperature, some organisms may be eliminated.
In dimorphic Malawian butterfly (Bicyclus anynana) in Africa, temperature also play

an important role. During the dry or cool season, the butterfly has mottled brown
characteristic that survives by hiding dead leaves on the forest floor. On the other hand,
during wet or hot season, where the butterfly routinely flies, the butterfly has a prominent
ventral eyespot that serves to deflect attacks from predators. This is made possible by the
gene Distal-less gene and the hormone 20-hydroxyecdysone.
Figure 62. Phenotypic plasticity in Malawian butterfly (Gilbert & Baressi, 2016)
Developmental Symbiosis
Symbiosis, an intimate relationship between organisms of different species, can also

influence development. This can occur in parasitism, when one partner benefits while
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causing harm to the other, and mutualism, in which both partners benefit from the relationship.
Typically, these relationships involve one organism that is larger than the other. The larger
organism is called the host, while the smaller one is called the symbiont. There are some
cases in which symbionts are tightly integrated into the host organism that the host cannot
develop without them. Holobiont refers to persistent symbionts. Endosymbiosis generally
refers to the situation in which one cell lives inside another cell.
Vertical and horizontal transmission. As stated, some host organisms’

development completely depends on the presence of symbionts. It is imperative that these
symbiotic relationships must maintain their partnerships over successive generations. This
can be achieved through vertical and horizontal transmission. Vertical transmission refers
to the transfer of symbionts through generations by the germ cells, usually the eggs. This is
the case for the bacterial genus Wolbachia and in some species of invertebrates. Wolbachia
resides the egg cytoplasm in invertebrates, providing necessary signals for the development
of the individuals produced by those eggs. In Drosophila, the Wolbachia host’s nurse cell
microtubule system and dynein motors to travel into the developing oocyte. In the oocyte, the
bacteria enter every cell, becoming endosymbionts. The Wolbachia can provide resistance
against resistance and can also aid in propagation by entering ovaries and oocytes. Females
infected with Wolbachia can produce four times more eggs than the uninfected ones.
Horizontal transmission refers to the transfer of symbiont to a newly born organism free
from any symbiont. An example of horizontal transmission are the symbiotic gut bacteria
found in most animals, even in humans. Mammalian gut bacteria are found to be critical in
blood vessel formation in the intestine and regulating stem cell proliferation. Newly born
infants acquire these symbiotic bacteria as they pass through the birth canal and in a
mother’s skin.
Obligate developmental mutualism. Obligate developmental mutualism occurs

when the species involved are interdependent to the point that neither can survive without
the other. Example for these are lichens, in which a fungal and algal species form a
relationship to create a new species. Another example is the Wolbachia and the wasp,
Asobaria tabida. If the Wolbachia are removed from the female germ plasm of the wasp
species, the ovaries undergo apoptosis, resulting to the absence or sterility of eggs.
Developmental symbiosis in the mammalian intestine. As mentioned, bacteria

aids in regulating gut development, especially in mammals. In an experiment with mice, it
was observed that bacteria can induce the production of fucosyl transferase enzymes of
mouse intestinal villi. Moreover, normally occurring gut bacteria can upregulate several
transcription of mouse genes, including colipase (for nutrient absorption), angionenin-4
(blood vessel formation) and Sprr2 (fortifies the extracellular matrices lining the intestine).
Bacteria can also help regulate the development of the immune and nervous system.
It was reported that intestinal microbes are important for the maturation of mammalian gut-
associated lymphoid tissue (GALT), a tissue that mediates mucosal immunity and oral
immune tolerance.
Changes in a Woman’s Gut During Pregnancy. Gut bacteria changes dramatically

in a pregnant human, allowing the woman to respond to hormonal status and adapt to
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physiological stress in carrying a fetus. During pregnancy, the normal microbiota is altered
resulting to weight gain with the progressive insensitivity to insulin.
Figure 63. Phenotypic plasticity in Malawian butterfly (Gilbert & Baressi, 2016)
References
Associates Inc.
Self Help.
Matsumoto, T., & Yamaoka, Y. (Eds.). (2019). Microbiota: Current research and emerging
trends. ProQuest Ebook Central https://ebookcentral.proquest.com
Gilbert, L. I. (Ed.). (2011). Insect endocrinology. ProQuest Ebook

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Let’s Check!
Activity 15. Now, that you know some basic concepts about development and the
environment, let us try to check your understanding of these topics. Give what is asked below.
___________1. This is the ability to modulate development in the presence of predators.
___________2. These are chemicals secreted by predators that can induce defenses in its
prey.
___________3. This refers to discontinuous phenotypes elicited by the environment.
___________4. This describes the genomic encoding of potential phenotypes that can be
determined later in life by environmental factors.
___________5. This are different phenotypes produced by different environmental

conditions.
___________6. This is a protein responsible for the development of a larvae to a queen in
ants, wasps and bees species.
___________7. This is the smaller organism within a symbiotic relationship.
___________8. This is the transfer of symbionts from one generation to another through the
germ cells.
___________9. This is a type of symbiotic relationship in which both partners cannot survive
without the other partner.
___________10. This the larger organism within a symbiotic relationship.
Let’s Analyze!
Activity 15. Now, that you know some basic concepts about development and the
environment, it is important that you will be able to relate this in your future profession. I will
1. Aside from what is given above, what are other examples of reaction norm?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Aside from what is given above, what are other examples of polyphenism?
__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
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3. How can diet affect DNA methylation in honeybees?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What are other effects of altered microbiota in pregnant women?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. What role does a mother’s milk play for the baby’s developing gut microbiota?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 15. Based on your learnings about some basic concepts of development and the
environment, feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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1.
2.
3.
4.
5.
Keywords Index
Phenotypic plasticity Symbiosis
Developmental plasticity Vertical & Horizontal Transmission

Ecomorphs Symbionts
Kairomones Hosts

ULO-c. Discuss development and evolution, including developmental
mechanisms of evolutionary change.
Metalanguage

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1. Evolutionary developmental biology, or evo-devo is a new scientific field that

integrates developmental biology, paleontology, and population genetics that aim
to explain and define the diversity of life.
2. Evolution is the process of which variety of organisms are thought to have
developed and diversified from earlier forms during the history of the earth.
3. Descent with modification refers to the idea that species change over time and
may give rise to new species. It also includes the idea that species share a
common ancestor.
4. Developmental module is a set of biologically functioning systems which are
more intrinsically connected than extrinsically. Examples of developmental
module are morphogenetic fields, signal transduction pathways, imaginal discs,
cell lineages, insect parasegments and vertebrate organ rudiments.
5. Disassociation is the ability of one module to develop into other modules.
6. Molecular parsimony refers to the uniformity of molecules used in development
throughout different lineages across time.
7. Paralogues are members of a gene family homologous within a species.
8. Orthologues are genes that are homologous between species.
9. Deep homology refers to homologous pathways made of homologous
components used for the same function in both protostomes and deuterostomes.
10. Heterotropy is the spatial alteration of gene expression, which permits different
cells to take a new identity or to activate or inhibit a paracrine factor-mediated
process in a new area of the body.
11. Heterochrony is the shift in the relative order or timing of two developmental
processes.
12. Heterometry is the change in the amount of a gene structure or product.
13. Heterotropy affects the coding regions of the gene, changing the functional
properties of the protein to be synthesized.
Key Knowledge
Evolutionary developmental biology, or evo-devo is a new scientific field that

integrates developmental biology, paleontology, and population genetics that aim to explain
and define the diversity of life. This new facet in evolution and development studies is
anchored to the concept that development is the changes in gene expression and cell
position over time and change in development over time can be considered as evolution.
Preconditions for evolution, deep homology, and mechanisms for evolutionary change will
be discussed in this section.
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Preconditions for Evolution
The developmental structure of a genome is a precondition for evolution. Specifically,

large morphological changes that can arise during development is attributed to two conditions
that underlie the development of all multicellular organisms: modularity and molecular
parsimony.
Modularity. Developmental module is a set of biologically functioning systems which

are more intrinsically connected than extrinsically. This connection is said to be constant
across some clades and can play specific causal role in development. Examples of
developmental module are morphogenetic fields, signal transduction pathways, imaginal
discs, cell lineages, insect parasegments and vertebrate organ rudiments. Disassociation,
the ability of one module to develop into other modules, can cause divergence in organisms.
Since DNA regions that can form the enhancer genes are modular, any mutation to this gene
can cause deviations in the place and timing of gene expressions affected by the enhancer
gene. This in return, can cause different anatomical and physiological morphologies,
resulting to new selectable variation for natural selection to work on.
Molecular parsimony. Molecular parsimony refers to the uniformity of molecules

used in development throughout different lineages across time. Examples are transcription
factors (ex. BMP, Hox, and Pax groups), paracrine factors, adhesion molecules and
transduction cascades. These molecules serve as a ‘small toolkit’ for development. Genes
like Hox gene, the globin genes, the collagen genes, Distal-less genes, and many paracrine
factor families are considered as paralogues, or homologous genes that have similarities
because they are descents from a common ancestor and are not the product of convergence
for a particular function. The original gene are considered to have duplicated and there are
subsequent independent mutations of the original duplicated gene, creating the mentioned
gene families related by common descent. Orthologues, on the other hands, are genes that
are homologous between species, as opposed to paralogues where members of a gene
family being homologous with a species.
Deep Homology
Deep homology refers to homologous pathways made of homologous components

used for the same function in both protostomes and deuterostomes. An evidence for deep
homology is the conserved similarities and functions of these pathways and components
even with phylogenetic divergence of the organisms over millions of years. An example for
this the same set of instruction that forms the nervous system of both fruit fly (protostomes)
and mouse (deuterostomes) shown below.
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Figure 64. Same Set of Instruction for the Nervous System in the Protostome, Fruit
Fly, and the Deuterostome, Moise (Gilbert & Baressi, 2016)
Mechanisms of Evolutionary Change
Changes between species, such as in humans and chimpanzees, were mostly

attributed to where, when, and how much gene are activated, rather than the allelic
substitutions of the genes that encode protein sequences. Wallace Arthur (2004) established
four ways in which changes at the level of gene expression can occur to generate phenotypic
variation for natural selection to work: (1) heterotropy (change in location), (2) heterochrony
(change in time), (3) heterometry (change in amount), and (4) heterotypy (change in kind).
Heterotropy. Heterotropy is the spatial alteration of gene expression, which permits

different cells to take a new identity or to activate or inhibit a paracrine factor-mediated
process in a new area of the body. Heterotropy is demonstrated in the development of bat
wings. Webbing of the forelimbs in bats is caused by the same blocking action of the BMPs
in duck hindlimb webbing. Unlike other mammals, bats express Fgf8 (a component of FGF
signaling), to promote webbing. If FGF signaling is stopped, apoptosis of the forelimb
webbing can occur.
Heterochrony. Heterochrony is the shift in the relative order or timing of two

developmental processes. Examples of heterochrony in vertebrates include the extend
growth of the human brain and the heterochronies of metamorphosis in amphibians.
Heterometry. Heterometry is the change in the amount of a gene structure or product.

An example of this is the observed difference of the evolution of the blind Mexican cavefish.
It was noted that the overproduction of the Sonic hedgehog protein (Shh) in the midline
prechordal plate downregulates the Pax6 gene, thus inhibiting eye formation.
Overexpression of the Shh gene also caused the cavefish’s jaw size and number of taste
buds to increase.
Heterotypy. Heterotropy affects the coding regions of the gene, changing the
functional properties of the protein to be synthesized and can have a profound effect in both
animal and plant evolution.
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Developmental Constraints on Evolution
Developmental constraint is the collective term used to refer to the limited number
and forms of possible phenotypes across the animal kingdom. This is because of the limited
interactions that are possible among molecules and between modules. There are three major
categories of developmental constraints: physical, morphogenetic, and pleiotropic
constraints. Physical constraints may include the laws of diffusion, hydraulics, and physical
support. Morphogenetic constraints may include the reaction-diffusion mechanisms. Lastly,
pleiotropic constraints refer to the constraints caused by pleiotropy, or the ability of a gene to
play different roles in different cells. When genes acquire new function, their activities may
extend to other module making evolutionary change difficult.
Selected Epigenetic Variation
Some variations produced by the environment can be inherited and selectable. This
is made possible by environmental agents that might change the germline DNA, rather than
somatic DNA alone. There are three process in which environmental induced changes can
be transmitted from generation to generation: epialleles, symbionts, and genetic
assimilation.
Epialleles. Epialleles are variants of chromatin structure that can be inherited

between generations. Epialleles are variations in DNA methylation patterning that may affect
the germ line of and organism, thus, transmissible to the offspring. Examples of epiallelic
inheritance may include diet-induced DNA methylation, endocrine disruptor-induced
DNA methylation, and behavior-induced DNA methylation. A general example of
variations in epigenetic forms caused by epialleles with difference in DNA methylation
patterning can be observed among toadflax shown below.
Figure 65. Epigenetic forms of toadflax. (A) Typical Linaria, with unmethylated
cyloidea gene; (B) Linaria, with heavily methylated cyloidea gene (Gilbert & Baressi,
2016)
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Symbionts. Contributing to phenotypic plasticity on an organism, symbionts can also

pass on this benefit to an organism’s offspring via the germ line. With this, symbionts can be
sources of selectable variation. An example of this is the community of bacteria in the pea
aphid Acrythisiphon pisum, that provide different phenotypic traits for the aphid. For instance,
pea aphids with the bacteria species Ricketsiella often give the adult pea aphids a red color.
On the other hand, pea aphids without Ricketsiella become green adult pea aphids.
Figure 66. Difference in the color of adult pea aphids in terms of the presence of the
bacteria Ricketsiella. (A) Without Ricketsiella turns adult pea aphid red; (B) With
Ricketsiella turns adult pea aphid green (Gilbert & Baressi, 2016)
Genetic Assimilation. Genetic assimilation refers to the continuous selection of

environmentally induced traits to stabilized it genetically and become heritable. The
phenotypic trait is initially produced in response to some environmental influence, and
through a process of selection, the phenotype is taken over by the genotype so that the
phenotype still forms even with the absence of the environmental influence.
References
Associates Inc.
162
Phone No.: (082)300-5456/305-0647 Local 134
Self Help.
Matsumoto, T., & Yamaoka, Y. (Eds.). (2019). Microbiota: Current research and emerging
trends. ProQuest Ebook Central https://ebookcentral.proquest.com
Let’s Check!
Activity 16. Now, that you know some basic concepts about development and evolution, let
us try to check your understanding of these topics. Give what is asked below.
___________1. This is the shift in the relative order or timing of two developmental processes.
___________2. This refers to the homologous pathways and components that spell for same
function in both protostomes and deuterostomes.
___________3. Genes that are homologous between species.
___________4. Genes that are homologous within species.

___________5. This refers to the ability of one module to develop differently from other
modules.
___________6. This refers to the concept of unity of type and conditions of existence by
Charles Darwin.
___________7. This refers to the spatial alteration of gene expression.
___________8. This refers to the change in amount of a gene product or structure.

___________9. This refers to the change in the coding region of the gene.
___________10. This is a developmental constraint that includes the laws of diffusion,
hydraulics, and physical support.
Let’s Analyze!
Activity 16. Now, that you know some basic concepts about development and 163volution, it
is important that you will be able to relate this in your future profession. I will require you to
give what are asked:
1. What is the role of heterometry on the variations of Charles Darwin’s finches from
Galapagos and Cocos islands?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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2. What is the role of heterotypy in the evolution of pregnancy in mammals?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. Give an example of genetic assimilation in the natural environment.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What is/are the condition/s for symbionts to be inherited by the offspring?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. What is the significance of genetic assimilation?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 16. Based on your learnings about some basic concepts of development and
evolution, feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
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5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
Keywords Index
Evo-devo Dissociation
Modularity Paralogues
Molecular Parsimony Orthologues
Evolution Deep Homology
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Big Picture
PLANT DEVELOPMENT

a. Discuss plant life cycles, pollination, and fertilization.
b. Discuss embryogenesis, dormancy, germination, and vegetative growth in plants.
c. Discuss vegetative-reproductive transition, and senescence in plants.

ULO a. Discuss plant life cycles, pollination, and fertilization.
.
Metalanguage

1. Alternation of generation in plants alternate generations means that plants

have a multicellular haploid and multicellular diploid stages throughout their life
cycle.
2. Heterosporous plants make two distinct spores that develop into the male and
female gametophytes.
3. Homosporous plants have sporophytes which only produce one type of spore
within a structure called the sporangium.
4. Interspecific incompatibility ensures that pollen from one species will not
germinate or grow on the stigma of another plant species. Intraspecific
incompatibility occurs within species.
Key Knowledge
Plants have different development as compared to animals. Land plants have

originated from freshwater green algae and their transition to the land requires evolution that
protects the embryo. The following items below shows the differences between animal and
plant development.
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1. Plants do not gastrulate. Plants do not rely on gastrulation to establish its basic
tissue systems.
2. Plants have sporic meiosis. Unlike animals with gametes, plants have spores
produced by meiosis. The gametes are produced by mitotic divisions following
meiosis.
3. Plants’ life cycles have both diploid and haploid multicellular stages referred to
as alternation of generations.
4. Germ cells are not set aside early in development.
5. Plants undergo extended morphogenesis. Example of this are the meristems that
continues to grow even after the plant’s maturity.
6. Plants have effective developmental plasticity which allows them to effectively
adapt to their surroundings especially that they are immotile.
7. Plants can tolerate higher genetic loads compared to animals. Plant phenotypes
do not easily change with mutations in the genome.
Plant Life Cycle
Plants alternate generations. This means that plants have a multicellular haploid and
multicellular diploid stages throughout their life cycle. Gametes develop in the multicellular
gametophyte. Fertilization will give rise to a multicellular diploid sporophyte, which produces
haploid spores via meiosis. This type of life is called haplodiplontic life cycle. On the other
hand, animals have diplontic life cycle, in which only the gametes are in the haploid state.
Mosses and other nonvascular plants such as liverworts and hornworts have life
cycles dominated by gametophytes, which means that they are usually larger and longer
living than the sporophytes.
Mosses are considered as heterosporous, in which they make two distinct spores
that develop into the male and female gametophytes. Male gametophytes develop
antheridia, male reproductive structures that produce sperm by mitosis, while female
gametophytes develop archegonia, the female reproductive structure that produce the eggs
by mitosis. Typically, sperms travel to a neighboring plant, via water droplets and are
chemically attracted to the archegonium, resulting to fertilization. The diploid embryonic
sporophyte develops within the archegonium and the mature sporophyte remains attached
to the gametophyte. The gametophyte nourishes both the embryo and the mature sporophyte
since they are not photosynthetic. Meiosis within the capsule of the sporophyte produces
haploid spores that are released and will germinate to a male or female gametophyte.
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Figure 67. Life Cycle of Moss (Urry et al., 2017)
Ferns, on the other hand, are considered as homosporous, that is the sporophyte
only produces one type of spore within a structure called the sporangium. One gametophyte
can produce both male and female sex organs. The greatest contrast between the mosses
and the ferns is that both the gametophyte and the sporophyte of the fern photosynthesize
and are thus autotrophic; the shift to a dominant sporophyte generation is taking place.
Figure 67. Life Cycle of Fern (Urry et al., 2017)

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Figure 68. Gametophyte-sporophyte relationships in different plant groups (Urry et

al., 2017)
Gamete Production in Angiosperm
The gametophyte generation produces the gametes in angiosperms. Typically,

angiosperm gametophytes are associated with flowers. The gametes they produce join to
form the sporophyte, which is what is commonly seen as the plant body.
Figure 68. Parts of a Flower (Urry et al., 2017)

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The shoot meristem of the sporophyte produces a series of vegetative structures. At

a certain point in development, internal and external signals trigger a shift from vegetative to
reproductive (flower-producing) development, initiating the development of floral parts
modified from leaves. Sepals and petals are sterile organs originated from the first and
second whorls of the leaves. The stamen, the pollen producer, originates from the third whorl
of the flower, while the carpel, which originates, from the fourth whirl contains the female
gametophyte. The stamen has four groups of cells called the microsporangia or pollen sacs
within an anther. The microsporangia undergo meiosis to create microspores. The inner
wall of the pollen sac, the tapetum, provides nourishment for the developing pollen.
Source: https://s3-us-west-2.amazonaws.com/courses-images/wp-content/uploads/sites/1223/2017/01/31224613/Figure_32_01_02.png
Figure 69. Male and Female Reproductive Parts in Flower
Pollen
Pollen is a simple multicellular structure that carries the sperm. Its outer wall, the
exine, is composed of resistant material provided by both the tapetum (sporophyte
generation) and the microspore (gametophyte generation). On the other hand, intine, the
inner wall, is produced by the microspore. A mature pollen grain consists of two cells, one
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within the other. The tube cell contains a generative cell within it. The generative cell divides
to produce two sperm. The tube cell nucleus guides pollen germination and the growth of
the pollen tube after the pollen lands on the stigma of a female gametophyte. One of the two
sperm will fuse with the egg cell to produce the next sporophyte generation. The second
sperm will participate in the formation of the endosperm, a structure that provides
nourishment for the embryo.
Source: https://media-mycbseguide.s3.amazonaws.com/images/quesbank/12/bio/ch01/63261_r1.png
Figure 70. Parts of a Pollen
The Ovary
The carpel is formed from the fourth whorl of organs within the flower. It gives rise to
the female gametophyte and consists of the stigma, the style, and the ovary. In
angiosperms, the ovary wall develops into a fruit succeeding fertilization. Inside the ovary are
one or more ovules attached by a placenta to the ovary wall. Seeds are fully developed
ovules. Ovules has one or two outer layers of cells called the integument. The integuments
enclose the megasporangium, which houses the sporophyte cells that undergo meiosis to
create megaspores. The micropyle is a small opening in the integuments where the pollen
tube will grow. When the mature embryo disperses from the parent plant, the embryo is
accompanied by diploid sporophyte tissue in the form of the seed coat and the fruit.
Meiosis and unequal cytokinesis within the ovule often yield four megaspores. The
largest of the megaspores undergoes three mitotic division, producing a seven-celled
embryo sac containing eight nuclei. Two of these are called synergid cells that surrounds
the egg. One of these cells is the actual egg. The central cell, which contains two or more
polar nuclei, will fuse with the second sperm nucleus to develop into the polyploid endosperm.
There are three antipodal cells at the opposite end of the embryo sac from the synergids.
These cells degenerate before or during embryonic development.
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Source: https://botanycompanion.files.wordpress.com/2015/01/angiosperm-ovule-copy.jpg
Figure 71. Flowering Plant Ovule
Pollination and Fertilization
Pollination involves the interaction between the gametophytic generation of the pollen
(male) and the sporophytic generation of the stigmatic surface of the carpel (female).
Pollination can either involve within a single flower, or another flower. Perfect flowers
contain both and female gametophytes. Staminate flowers do not have carpel, while
carpellate flowers lack stamens. Plants that have both staminate and carpellate flowers,
such as maize, are called monoecious plants. Plants that have staminate and carpellate
flowers on separate plants are considered as dioecious. Interspecific incompatibility
ensures that pollen from one species will not germinate or grow on the stigma of another
plant species. Intraspecific incompatibility occurs within species. An example for this is
self-incompatibility between the pollen and the stigma of the same individual.
When the pollen and stigma are compatible, the pollen hydrates and the pollen tube
emerge. The pollen tube grows down the style of the carpel toward the micropyle, carrying
the tube nucleus and the sperm cells at the growing tip by bands of callose, a complex
carbohydrate. Calcium is known to contribute to the growth of the pollen tube. Calcium
usually accumulates in the tip of the pollen tubes, where open calcium channels are plenty.
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Sources: https://www.toppr.com/ask/content/story/amp/fertilization-in-plants-51976/
Figure 72. Germinating Pollen Tube
The growing pollen tube enters the embryo sac through the micropyle, growing
through one of the synergids. As the two sperm cells are released, double fertilization occurs
where one sperm cell fuses with the egg, producing the zygote which will develop into the
sporophyte, and the second sperm cell fuses with the bi- or multinucleate central cell, giving
rise to the endosperm. The endosperm nourishes the growing embryo.
Figure 73. Gametophyte Development in Angiosperm (Leyser & Day, 2002)

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References
Leyser, O., & Day, S. (2002). Mechanisms in plant development. ProQuest Ebook
Gilbert, S. (2003). Developmental biology (7th Ed.). Sinauer Associates Inc.: USA
Self Help.
Plant development and evolution. (2019). ProQuest Ebook

Let’s Check!
Activity 17. Now, that you know some basic concepts about plant life cycle, pollination, and
fertilization, let us try to check your understanding of these topics. Identify what is asked
below:
___________1. Attaches ovules to the ovary wall.
___________2. These encloses the megasporangium.
___________3. The cell that divides to produce two sperms in a pollen.
___________4. This is the structure that produce eggs in mosses.
___________5. Plants that can make two types of spores.
___________6. Plants that can make only one type of spores.
___________7. The fourth whorl of organs modified from leaves in flowers.
___________8. These cells undergo meiotic division to produce microspores in stamens.

___________9. The first and second whorl of organs modified from leaves in flowers.
___________10. The opening in the integuments where the pollen tube will grow.
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Let’s Analyze!
1. What are the differences in pollination and fertilization between angiosperms and
gymnosperms?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Give other examples of monoecious and dioecious plant species. Is there any
evolutionary significance to this characteristic?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. What is the function of the synergid and antipodal cells within the embryo sac?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What is the significance of self-incompatibility during pollination?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. What is a S locus? What is its role in self-incompatibility in plants?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 17. Based on your learnings about some basic concepts about plant life cycle,
pollination, and fertilization, feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
Keywords Index
Plant Life Cycle Gametophyte
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Sporophyte Pollination
Fertilization Carpel
Stamen Endosperm

ULO-b. Discuss embryogenesis, dormancy, germination, and
vegetative growth in plants.
Metalanguage

1. Embryogenesis, in plants, covers development from the time of fertilization until

dormancy occurs.
2. Dormancy in seed can be defined as the state or conditions where the seeds
are restricted to germinate until certain environmental cues causes them to break
dormancy.
3. Germination refers to the process by which an organism grows from a seed or
a spore.
4. Vegetative growth in plants refer to the development of plants between
germination and flowering. During this stage extensive photosynthesis and
accumulation of resources need for flowering and reproduction occur.
Key Knowledge
Embryogenesis
Succeeding fertilization is the process of embryogenesis. In plants, embryogenesis

includes the development from the time of fertilization until dormancy occurs. Majority of plant
species have their zygotes divided in a plane perpendicular to the long axis of the embryo
sac to produce a large basal cell close to the micropyle, and a small terminal cell near to
what was the central cell and is now the developing triploid endosperm. The first distinction
between cells during embryogenesis shows the cells which will form the main body of the
embryo and those that will produce the suspensor, which becomes evident after cell division.
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Suspensor is a filament that connects the embryo to the maternal tissue close to the
micropyle.
During the first stages of plant embryogenesis, the embryo undergoes cell divisions
with little or no increase in the embryo’s total size. This results to a ball of cells called the
globular embryo. The three tissues in plants - dermal, vascular, and ground tissue - are
established in this stage. The epidermis forms from the outermost layer of cells. They are
typically smaller as compared to the cells of the underlying tissues. Protoderm often refers
to the epidermis of the embryo and their immature regions of growing shoots and roots. The
vascular tissue during this stage is characterized by large cells with abundant vacuoles,
and a strand of smaller, less vacuolated cells. In this immature state, it is called the
procambium. Lastly, the larger cells between the epidermal and vascular tissue forms the
ground tissue of the embryo. In this immature state, this is called the ground meristem. The
apical meristem, which are the precursors of the root and shoot, appear during or later in
embryogenesis. It is characterized by clusters of densely cytoplasmic cells at either end of
the procambial strand. Apical meristems give rise to all post-embryonic cell lineages. The
root apical meristem forms at the end of the procambial strand nearest to the suspensor,
while the shoot apical meristem develops far from the suspensor at the other end of the
procambium.
The formation of the cotyledon near the site of the shoot apical meristem signifies the
end of the globular stage of embryo development. In dicots, two cotyledons develop,
changing the globular embryo to a heart-shaped embryo. The heart-shaped embryo
elongates to form the torpedo-stage embryo, while retaining the same pattern of tissues and
organs. Monocots, on the other hand, form a single cotyledon.
Figure 73. Embryogenesis in Arabidopsis thaliana (Leyser & Day, 2002)
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In angiosperms, growth often pauses between the end of embryogenesis and the
start of germination, but the extent of embryo development before this pause varies between
species. In some species, their embryos have limited development, and a shoot apical
meristem and a root apical meristem form but immediately become dormant. In some species,
growth occurs at both apical meristems before dormancy. The shoot apical meristem forms
the plumule, while the root apical meristem grows a short length of root called the radicle.
In grasses, the coleoptile and the coleorhiza enclose the plumule and radicle, respectively.
Source: https://www.drgpbiology.com/dicot-and-monocot-embryo/
Figure 74. Dicot and Grass Embryo
Dormancy
Dormancy ends embryogenesis. An embryo prepares for dormancy by accumulating

storage carbohydrates, proteins, and lipids. Eventually, the high level of zygotic gene
expression and fast metabolism at the early stages of embryogenesis slow down and the
connection of the seed to the ovary is severed by the degeneration of the adjacent supporting
sporophyte cells. Seed dormancy can last for weeks or years. The hormones abscisic acid
and gibberellin are necessary in dormancy. The abscisic acid maintains dormancy in many
species while gibberellins are essential in breaking dormancy.
Germination
Germination signifies the postembryonic phase of plant development. During

germination, extensive cell elongation forces the root outside the seed case, carrying the
shoot upwards. This process requires the optimal combination of external cues such as
moisture, temperature, oxygen, and light.
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In general, dormant seeds are imbibed, or added with water to let the seed rehydrate,
prior to the germination process. During imbibition, the seed soaks up with large volumes
of water, swelling to many times than its original size. Most of the germination plant draws
on the nutrient reserves in the endosperm or cotyledons. The breakdown of starch in the
sugar are made possible using the hormone gibberellin. The primary embryonic root, radicle,
emerges primarily from the seed to enhance water uptake. As the shoot reaches the surface,
light triggers the differentiation of chloroplasts. The delicate shoot tip is protected as it pushes
through the soil by the cotyledon, the bending of the epicotyl, and the coleoptile in monocots.
There are two types of seedlings distinguished by their germination. If the elongation
occurs between the cotyledons and the radicle (in the hypocotyl), the cotyledons are lifted
above the ground, making the seedling as epigeal. On the other hand, if elongation occurs
between the cotyledons and the shoot apical meristem (in the epicotyl), the cotyledons stay
underground, and the seedling is called hypogeal. In both cases, the germinating shoot
grows upward in hook shape so that it pulls rather than pushes the shoot tip through the soil.
Figure 75. Epigeal and Hypogeal Germination (Leyser & Day, 2002)
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Vegetative Growth in Plants
Meristems
Cells in the meristems are analogous to the stem cells in animals. Through mitotic
division, the meristems give rise to one daughter cell that continues to be meristematic and
other than differentiates. There are three categories of meristems: apical, lateral and
intercalary.
Apical meristem. As mentioned, apical meristems occur at the developing shoot

and root tips. Specifically, root apical meristems (RAM) produce the root cap consisting of
lubricate cells that are sloughed off as the meristem is pushed through the soil by cell division
and elongation of the proximal cells. The three-tissue system of the root also arise from the
root apical meristem. From the circumference inwards, the main tissues are the epidermis,
the cortex, the endodermis, the pericycle, and the central core of vascular tissue. New
root apical meristems are derived from the tissue within the core of the root and emerge
through the ground tissue and dermal tissue. Root meristems can also be derived secondarily
from the stem of the plants. Shoot apical meristem (SAM) produces the stems, leaves ad
reproductive structures. The axillary shoot apical meristem derived from SAM forms in the
axils, or the angles between the stem and leaf. The size of SAM is controlled by intercellular
signals between the layers of the meristems.
Lateral and intercalary meristem. These are cylindrical meristems located in

shoots and roots that usually responsible for secondary growth in plants. Monocot stems lack
lateral meristems, but often have intercalary meristems inserted in the stems between
mature tissues.
Root Development
In roots, both radial and axial patterning starts during embryogenesis and continues
throughout development as the primary root grows and lateral roots emerge from the
pericycle cells deep within the roots. Most of the cells in roots are plastic and their position is
the primary determinant of fate in early root development. In wild-type Arabidopsis, there are
two layers of root ground tissue, where the outer later becomes the cortex and the inner later
becomes the endodermis. The genes involved in the formation of the two layers are the
SCARECROW (SCR) and SHORT-ROOT (SHR) genes. Specifically, SCR gene is essential
for the asymmetric cell division in the initial layer of cells, creating a larger cortex cell, and
smaller endodermal cells, while SHR genes is responsible for endodermal cell specification.
Moreover, both genes are essential for normal gravitropic response, which is dependent on
normal endodermis formation. Furthermore, plant hormones also influence root development.
For instance, distribution of auxin organizes the axial pattern.
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Shoot Development
Shoot meristems are the origins of aboveground architectures of different plant

species. The amount of axillary bud outgrowth affects shoot architecture. The shoot tip
regulates the branching pattern, called the apical dominance, which is highly influenced by
plant hormones. The hormone auxin is produced by young leaves and transported toward
the base of the leaf, where it can suppress the outgrowth of axillary buds. Cytokinin can also
release buds from apical dominance. Branching can be regulated by both environmental
signals and genes.
Leaf primordia, or cluster of cells fated to become leaves, are derived at the
periphery of the shoot meristem. The node is the union of a leaf and the stem, while the stem
tissue between nodes is called an internode. Phyllotaxy, or the positioning of leaves on the
stem involves the communication among existing and newly formed leaf primordia.
Leaf Development
Leaf developments include the scope of commitment to becoming a leaf, establishing

the leaf axes and morphogenesis. In some research on ferns and angiosperm, the youngest
leaf primordia are not yet determined to make a leaf. This leaf primordia can still develop as
shoots in culture. The programming for leaf development occurs later. The radial symmetry
of the leaf primordium becomes dorsal-ventral, or flattened, in all leaves. The proximal-distal
and lateral are also established. The distinguishing shapes of leaves result from regulation
of cell division and cell expansion as the leaf blade develops. Aside from differential cell
growth, apoptosis can also be involved in the shaping of a leaf.
Generally, there are two categories of leaves: the simple and compound.
Compound leaves are composed of individual leaflets while simple leaves have a single leaf
blade. There are several perspectives as to the mechanism of development in these leaves.
For instance, compound leaves are highly lobed simple leaves, or modified shoots. Some
genes involved in leaf composition is the Class I KNOX genes involving STM and KNOTTED
1 (KN1) gene (observed in maize), and LEAFY gene observed in Arabidopsis and
snapdragon (which is called FLORICAULA gene).
References
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Self Help.

Let’s Check!
Activity 18. Now, that you know some basic concepts about embryogenesis, dormancy,
germination, and vegetative growth in plants, let us try to check your understanding of these
topics. Match the stage of plant development related to the items given below:
____1. Leaf primordia a. Embryogenesis
____2. Global embryo b. Dormancy
____3. Pause in seed metabolism c. Germination

____4. Imbibition d. Vegetative Growth
____5. Phyllotaxy
____6. Auxin
____7. Gibberellin
____8. Abscisic Acid
____9. Meristem
____10. Cotyledon
Let’s Analyze!
1. What is the evolutionary significance of seed dormancy?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Give the different plant hormones and their functions.
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__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. What is primary and secondary vegetative growth? What is its significance?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. What is the role of imbibition in the process of germination?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. Why are most seeds desiccated during dormancy?

__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 18. Based on your learnings about some basic concepts about plant life cycle,
pollination, and fertilization, feel free to write your arguments or lessons learned below:
1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
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__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
Keywords Index
Embryogenesis Dormancy
Germination Cotyledons
Vegetative growth Meristems
Leaf primordia Imbibition
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ULO-c. Discuss the vegetative-to-reproductive transition and
senescence in plants.
Metalanguage

1. Photoperiods refers to day lengths.

2. Phytochrome is a blue-green pigment in plants that regulates gene expression.
3. Vernalization refers to the cooling of seed during germination to accelerate
flowering when planted.
Key Knowledge
Vegetative-to-Reproductive Transition in Plants
Both vegetative and reproductive structures of the shoot are derived from the shoor
meristem formed during embryogenesis. In general, the vegetative-to-reproductive transition
in plants refers to the process of flowering. The timing of flowering is essential in ensuring
maximal reproductive successes. The transition begins with the signal form the leaves
moving towards the shoot apex to induce flowering, and oftentimes, these developmental
pathways leading to flowering at numerous control points different plant organs.
In woody perennials, a juvenile phase exists in which the plant cannot produce
reproductive structure even if all optimum environmental signals are present. The juvenile
plant can transition to the adult stage if the leaves or meristem acquire competence in
response to internal or external signals. Some experiments involving grafting and organ
culture experiment established that leaves could produce a graft-transmissible substance
that can induce flowering. This signal is only produced under specific photoperiods, while
other species are day neutral and will flower under any photoperiod. Some leaves, however,
are not competent to perceive or pass on photoperiodic signal. Instead, the phytochrome
pigments transduce these signals from the external environment. The phytochrome structure
is modified by red- or far-red light, and these changes can initiate a cascade of events
resulting to the production of either floral promoter or inhibitor. Floral inhibitors can also be
found in leaves, cotyledon, and roots in some species. Other species of plants change the
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competence of their meristems in responding to flowering signals during development. In

vernalization, or a period of chilling, the competence of both leaves and shoots are
enhanced to perceive or produce a flowering signal.
In ancestral angiosperm, it is believed that the terminal flower directly formed from
the terminal shoot apex. In modern angiosperm, however, variety of flowering pattern can
occur including axillary buds that can produce flowers. The transition of a vegetative
meristem to an inflorescence meristem includes axillary meristems that can produce floral
organs, but do not directly produce floral parts itself. The inflorescence meristem is perceived
to arise through the action of a gene that suppresses terminal flower formation. Example of
this gene is the CENTRORADIALUS (CEN) gene in that suppresses the expression of the
FLORICAULA (FLO) gene in snapdragon.
After the inflorescence meristems are established, the specification of the actual floral
meristem is underway. In plants, the genes responsible for this specification are called the
floral meristem identity genes. In snapdragon, the upregulation of the FLO gene transitions
the inflorescence meristems to become a floral meristem. Furthermore, the floral meristem
identity genes trigger a cascade of gene expression that activates the region-specifying
(cadastral) genes, which aids in specifying pattern by activating the transcription of floral
organ identity genes.
Figure 76. The shoot apical, inflorescence and floral meristem
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Senescence in Plants
In some plant species, flowering and senescence are closely related. For instance,
some individual flower petal in some species undergo senescence after pollinated. Fruit
ripening is an example of organ senescence. In some plants, like in whole-plants,
senescence leads to the death of the entire sporophyte generation. Plants that flower once
a year and undergo senescence are called monocarpic plants, while those that can live
thousands of years are called polycarpic plants. In plant species, the removal of flowers
and fruits can delay senescence. Typically, nutrients are reallocated from other parts of the
plant to support the development of the next generation. This causes the reproductive
structures to have less nutrients and can lead to whole-plant senescence.
References
Self Help.

Let’s Check!
Activity 19. Now, that you know some basic concepts about vegetative-to-reproductive
transition and senescence in plants, let us try to check your understanding of these topics.
Give what is asked below:
___________1. Plants that undergo senescence after flowering once a year.
___________2. Plants that can live thousands of years.
___________3. These are genes responsible for specifying inflorescence meristems to
become floral meristems.
___________4. These are genes responsible for specifying the floral meristems to respective
organs.
___________5. Refers to the flowering in angiosperm.

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Let’s Analyze!
Activity 19. Now, that you know some basic concepts about vegetative-to-reproductive
transition and senescence in plants, it is important that you will be able to relate this in your
future profession. I will require you to give what are asked:
1. What are the factors the common signs that a plant is undergoing senescence?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. Death in monocarpic plants is often genetically programmed while in polycarpic

plants, death is by accident. How is this so?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. What are the common factors that can trigger senescence in some plants?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. Give other examples of floral meristem identity and floral organ identity genes.
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. What is a phytochrome? What role does it play in vegetal-to-reproductive transition

in plants?
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
In a Nutshell…
Activity 19. Based on your learnings about some basic concepts about vegetative-to-
reproductive transition and senescence in plants, feel free to write your arguments or lessons
learned below:
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1. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
2. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
3. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
4. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________
5. __________________________________________________________________
__________________________________________________________________
__________________________________________________________________
__________________________________________________________________

1.
2.
3.
4.
5.
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Keywords Index
Vegetative Polycarpic
Reproductive Monocarpic
Senescence Floral meristem
Flowering Inflorescence meristem
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social media conduct (OPM 21.15) and personnel discipline (OPM 21.11).
3. All students are likewise guided by professional conduct as learners in

attending OBD or DED courses. Any breach and violation shall be dealt
with properly under existing guidelines, specifically in Section 7 (Student
Discipline) in the Student Handbook.
4. Professional conduct refers to the embodiment and exercise of the

University’s Core Values, specifically in the adherence to intellectual
honesty and integrity; academic excellence by giving due diligence in
virtual class participation in all lectures and activities, as well as fidelity in
doing and submitting performance tasks and assignments; personal
discipline in complying with all deadlines; and observance of data privacy.
5. Plagiarism is a serious intellectual crime and shall be dealt with

accordingly. The University shall institute monitoring mechanisms online
to detect and penalize plagiarism.
191
Phone No.: (082)300-5456/305-0647 Local 134
6. All borrowed materials uploaded by the teachers/Course Facilitators shall

be properly acknowledged and cited; the teachers/Course Facilitators
shall be professionally and personally responsible for all the materials
uploaded in the online classes or published in SIM/SDL manuals.
7. Teachers/Course Facilitators shall devote time to handle OBD or DED

courses and shall honestly exercise due assessment of student
performance.
8. Teachers/Course Facilitators shall never engage in quarrels with

students online. While contentions intellectual discussions are allowed,
the teachers/Course Facilitators shall take the higher ground in facilitating
and moderating these discussions. Foul, lewd, vulgar and discriminatory
languages are absolutely prohibited.
9. Students shall independently and honestly take examinations and do

assignments, unless collaboration is clearly required or permitted.
Students shall not resort to dishonesty to improve the result of their
assessments (e.g. examinations, assignments).
10. Students shall not allow anyone else to access their personal LMS
account. Students shall not post or share their answers, assignment or
examinations to others to further academic fraudulence online.
11. By handling OBD or DED courses, teachers/Course Facilitators agree

and abide by all the provisions of the Online Code of Conduct, as well as
all the requirements and protocols in handling online courses.
12. By enrolling in OBD or DED courses, students agree and abide by all the
provisions of the Online Code of Conduct, as well as all the requirements
and protocols in handling online courses.
MONITORING OF OBD AND DED
1. The Deans, Asst. Deans, Discipline Chairs and Program Heads shall be responsible
in monitoring the conduct of their respective OBD classes through the Blackboard
LMS. The LMS monitoring protocols shall be followed, i.e. monitoring of the conduct
of Teacher Activities (Views and Posts) with generated utilization graphs and data.
192
Phone No.: (082)300-5456/305-0647 Local 134
Individual faculty PDF utilization reports shall be generated and consolidated by

program and by college.
2. The Academic Affairs and Academic Planning & Services shall monitor the conduct
of LMS sessions. The Academic Vice Presidents and the Deans shall collaborate to
conduct virtual CETA by randomly joining LMS classes to check and review online
the status and interaction of the faculty and the students.
3. For DED, the Deans and Program Heads shall come up with monitoring instruments,
taking into consideration how the programs go about the conduct of DED classes.
Consolidated reports shall be submitted to Academic Affairs for endorsement to the
Chief Operating Officer.
193

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College of Arts and Sciences Education

General Education – Science

Physically Distanced but Academically Engaged

Self-Instructional Manual (SIM) for Self-Directed Learning (SDL)

Name of Teacher: ______________________________

THIS SIM/SDL MANUAL IS A DRAFT VERSION ONLY; NOT FOR REPRODUCTION

Prepared by Jessa Mae B. Patalita

Week 1 to 3: DEVELOPMENTAL ANATOMY AND GENETICS 11

The Cycle of Life 12

Week 4 to 5: CELL-TO-CELL COMMUNICATION IN DEVELOPMENT 36

The Epithelial-Mesenchymal Transition 48

Week 6 to 7: FERTILIZATION AND EARLY DEVELOPMENT IN VERTEBRATES 53

Structure of the Gametes 54

Week 8 to 9: EARLY DEVELOPMENT IN VERTEBRATES AND LATE 76

Early Amphibian Development 76

Formation of Limb Buds 102

Week 12 to 13: POSTEMBRYONIC DEVELOPMENT 120

Amphibian Metamorphosis 121

Week 14 to 15: DEVELOPMENT IN HEALTH AND DISEASE, 141

Genetic Errors of Human Development 142

Week 16 to 18: PLANT DEVELOPMENT 166

Plant Life Cycle 167

Vegetative-to-Reproductive Transition in Plants 186

Course Outline: BIO 238/L – Developmental Biology

Course coordinator: Jessa Mae B. Patalita

Course Outline Policy

Areas of Concern Details

If disapproved by the course coordinator, you can

You can also meet the course coordinator in person

For students who have not created their student e-

You can also contact the course coordinator through

For students who have not created their student

CO: In Developmental Biology, we will discuss the development of plants and

Week 1 to 3: Unit Learning Outcomes (ULOs):

Big Picture in Focus

In this section, the essential terms relevant to Developmental Biology and to

1. Developmental Biology is the study of the embryonic and developmental process

The Cycle of Life

In general, all animal collectively undergoes the process of embryogenesis, the

Figure 1. Life Cycle of a Frog (Gilbert and Baressi, 2016)

4. Organogenesis occurs after gastrulation and mainly involves the formation of

The Life Cycle of Rana pipiens, the Leopard Frog

Figure 4. Organogenesis in Rana pipiens.

Figure 4. Organogenesis in Rana pipiens (Gilbert and Baressi, 2016)

The metamorphosis of a tadpole to an adult frog will be initiated by hormones from

Anatomic Approaches to Developmental Biology

Preformationist hypothesis was established by Albrecht von Haller and Charles

By 1820s, Christian Pander established the three germ layers – ectoderm,

Evolutionary change is often based on developmental change. Without changes in

believed that embryonic resemblance is a strong support for genetic connectedness. He

Before Darwin’s revolutionary work, some taxonomists classify organisms according

Figure 5. Analogous and homologous structures

Homology and analogy are two essential concepts in evolutionary embryology.

Medical Embryology and Teratology

Figure 6. Piebaldism in human and mouse

Abnormalities caused by exogenous agents such as chemicals, radiation, viruses or

Figure 7. Phocolemia in a human baby

Streelman, J. T., & Streelman, J. (Eds.). (2014). Advances in evolutionary developmental

_____ 6. Completion of the number of chromosomes f. Gametogenesis

_____ 10. Development of gametes

2. Give other examples of malformations and disruptions prominent in the Philippines.

4. What is the importance of Developmental Biology in the field of Taxonomy and