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History of Embryology

This document summarizes the history and key concepts in embryology. It describes the stages of development from zygote to embryo to fetus. It then outlines early theories from Plato and Aristotle and important contributions from scientists like William Harvey, Antoni van Leeuwenhoek, and Ernst Haeckel. It discusses the debate between epigenesis versus preformation and the later synthesis of embryology with genetics. Key concepts in model organisms, reproduction, gametogenesis, and the male and female reproductive systems are also summarized.
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0% found this document useful (0 votes)
48 views35 pages

History of Embryology

This document summarizes the history and key concepts in embryology. It describes the stages of development from zygote to embryo to fetus. It then outlines early theories from Plato and Aristotle and important contributions from scientists like William Harvey, Antoni van Leeuwenhoek, and Ernst Haeckel. It discusses the debate between epigenesis versus preformation and the later synthesis of embryology with genetics. Key concepts in model organisms, reproduction, gametogenesis, and the male and female reproductive systems are also summarized.
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© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
Available Formats
Download as PDF, TXT or read online on Scribd
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HISTORY OF EMBRYOLOGY

● Zygote – upon fertilization; sperm egg cell


● Embryo – stops being an embryo during gestational stage; 8​th​ week from the time
it was a zygote
● Fetus – 9​th​ week (humans);
● Gestational stage – counting from time of fertilization
● Pregnancy – counting before fertilization including ovulation; 2 weeks earlier
than fertilization
● marine -> amphibious -> terrestrial
● fertilization can be inner (marine and amphibious) or external (terrestrial, can
also be internal)
● amniotes – egg can develop in land (amniotic fluid)

Early History
1. Plato
- concept of souls
- vegetative soul brings about life (plants stop here)
- sensitive soul enables sensations (animals)
- spiritual soul enables thinking (humans)
2. Aristotle
- systematic observations of embryos
- diff stages of development of chick embryos
- multiple ways organisms reproduce

Renaissance History
1. William Harvey
- work on blood
- expanded and corrected Aristotle’s work
- epigenesis – egg developing into embryo
- omne vivum ex ovo – presence of ova even in humans
2. Antoni van Leeuwenhoek
- animalcules
- tiny preformed human was already present in these animalcules
- preformationist period – homunculus – malpighi and swammerdam were the
chief ovists
3. Lazzaro Spallanzani
- FIRST ARTIFICIAL INSEMINATION – frog eggs
Epigenesis and Vitalism
1. Caspar Friedrich Wolff
- morphogenesis
- development of structure out of structureless yolk material
2. Johann Friedrich Blumenbach
- vis vitalis - vital force, immaterial virtue or life force
- Vitalism – explained how structure could develop from an amorphous state
- serious attempts to discover the nature of this force
3. Carl Ernst von Baer
- made significant strides in descriptive embryology - VITAL FORCE
- many similarities between embryos of vertebrates particularly amniotes

Ontogeny recapitulates Phylogeny


● ontogeny – life history (juvenile to adult)
● phylogeny – evolutionary history

1. Ernst Haeckel
- leading authority in embryology (late 1800s)
- ontogeny recapitulates of phylogeny – individual development progresses
through adult stages of the organisms ancestors
- race and human development and evolution were later incorporated into the
pseudo-scientific basis for Nazism

Synthesis of Embryology with Genetics


1. AUGUST WEISMANN
- germ plasm theory
- 1892 self-reproducing determinants – guiding agents in morphogenesis
- determinants were located on some newly discovered structures: chromosomes
- cell differentiation – result of cells acquiring diff chromosomes during replication
(not during fertilization)
2. Oscar Hertwig and Richard Hertwig
- 1​st​ demonstration of fertilization – sea urchins
- existence of polar bodies – parts of the immature egg cells
- significant advances in understanding of meiosis

Experiment Embryology
1. Hans Spemann
- induction – result of organizer effect; a group of cells would influence how other
cells develop
- organizer cells – the ones that influence
- nobel prize for Medicine – ORGANIZER EFFECT – underlying role of the dorsal
lip of the blastopore

Model Organisms
- prior to 1900s
- no model organisms targeted by large numbers of scientists
- isolated advances – work of individual genius

Thomas hunt morgan – fruit fly


Wilhelm roux – frog
hans Driesch – sea urchin
REPRODUCTIVE SYSTEM
MALE REPRODUCTIVE SYSTEM

● Testis
- male sex gland located in the scrotum
- produces sperm and androgens
● Seminiferous tubules
- lobes/structural units found in the testis
- responsible for spermatogenesis - production of sperm
● Primordial/stem cells
- early stages
- spermatogonium
● Spermatozoa
- mature sperm cells
- has all the necessary components
- unable to move
- head, neck,tail
- it has to undergo intense training in the epididymis – will be activated,
learn how to move – spermiation
● Epididymis
- sperm storage and maturation
● Scrotum
- sac-like
- descends so that testis is farther away from the body when cold; ascends
when it is hot
● Vas deferens
- transports sperm from the scrotum to prostate gland
- delivers mature sperm cells to the seminal vesicles then to the prostate
gland
● Seminal vesicles
- secretes fructose-rich semene
- provide energy to the sperm cell
● Prostate gland
- contributes milky alkaline fluid that assists sperm activation
- semen is coagulated (semi-solid) - has difficulty moving, 30 mins to liquify
for swimming purposes
● Cowpers/ bulbourethral gland
- contributes mucus to semen
● Urethra
- exit
● Penis
- delivers sperm into the female genital tract

TESTIS HISTOLOGY

- spermatogonium – undergoes mitosis to produce more spermatogonia,


differentiate primary spermatocyte, duplicates its chromosomes and undergoes
1​st​ meiosis cycle, secondary spermatocyte, 2​nd​ meiosis – spermatids –
round/spherical cells, undergo spermatogenesis
- spermiogenesis – grow a tail and a head
Sertoli cells
- maintain close contact with spermatogenic cells to provide structural and
metabolic support
- a single cell extends from the basement membrane to the lumen of the
seminiferous tubule
Sertoli Cell Functions
- maintain the environment necessary for development and maturation via the
blood-testis barrier
- secretes substances initiating meiosis
- secrete supporting testicular fluid
- secrete androgen-binding protein, which concentrates testosterone in close
proximity to the developing gametes
- secrete hormones affecting pituitary gland control of spermatogenesis,
particularly the polypeptide, inhibin
- phagocytose residual cytoplasm left over spermiogenesis

FEMALE REPRODUCTIVE SYSTEM

● Ovary
- female sex gland that produces ovum
● Uterus
- where fetus develops
- role up to the cervix is to deliver/ passageway
● Oocytes/ovum/egg cells
- follows meiosis 1 and 2, only 1 daughter cell proceeds to the next phase,
the other one becomes a polar body which eventually degenerates
● Oogonium
- primary oocyte – when it splits, one becomes primary oocyte and the other
becomes a polar body which cannot undergo fertilization, it will become a
secondary oocyte by fertilization, follicular cells
Meiosis
- reduction division
- homologous recombination or “crossing over”
- 4 genetically different haploid daughter cells (23 chromosomes in humans)
primary oocyte
- surrounds itself with follicular cells (Sertoli cells) which further grows to become
the zona pellucida – space cavity (avum?)
- 1 oogonium - 1 ovum
secondary oocyte
- egg cell is arrested at metaphase (has 46 chromosomes) ; upon fertilization, it
will split to 23 chromosomes

Gametogenesis
- production of gametes (sperm and egg) at the end of Meiosis II
- “gameto” – gametes
- “genesis” – to create/ make

Gametogenesis phases:
1. multiplication
- primordial germ cells undergo mitosis
- production of oogonia and spermatogonia

2. growth (process of maturing ; pag naging spermatocyte, oocyte, secondary, etc.)


3. maturation/ differentiation (when spermatids become sperm cells)
- primary spermatocyte -> 2 secondary spermatocytes -> 4 spermatids
which undergo differentiation (spermatogenesis)
- primary oocyte
Sperm structure: Acrosome
- golgi apparatus becomes the acrosome (responsible for dissolving cells
surrounding the ovum in order for the sperm cell to penetrate)
- golgi is at the north pole
- mitochondria moves to the south pole (the next middle ; becomes the neck) to
produce energy for the flagellum extending outside the cell
- derived from golgi apparatus and contains hydrolytic enzymes released when
sperm reaches an ovum, digesting the outer membrane of the egg and allowing
penetration of the sperm
Sperm structure: Midpiece
- contains numerous mitochondria that provide energy for movement of the tail

1. spermatogenesis (production of sperm cells)


- spermiogenesis (grow head and tail)
[ ] golgi becomes acrosome
[ ] mitochondria moves to posterior (neck)
[ ] centrioles move out of the spermatid ; becomes the tail
- spermiation (activation of the sperm cells in the epididymis thru vas
deferens)
- occurs upon release of mature spermatozoa from the protective
sertoli cells into the lumen of the seminiferous tubules
- removes remaining unnecessary cytoplasm and organelles
- mature but non-motile reptile are transported to the epididymis,
where in they acquire motility and become capable of fertilization
- gonads: testes/ testis
- continual production of spermatogonia
- spermatogonium - males, lifetime, can undergo replication

2. oogenesis (production of egg cells)


- gonads: ovaries/ ovary
- has 2 million primary oocytes in her ovaries
- by age 7, 300,000 remain – the rest are reabsorbed
- 400 to 500 oocytes will be released during the reproductive years
- oocytes remain arrested at prophase until ovulation
- at the time of ovulation, oocytes proceed up to Metaphase II
- penetration of sperm induces the secondary oocyte and the first polar body
to complete meiosis II
- if unfertilized, oocyte degenerates
- limited number of oogonium
- oogonium
- upon birth - predetermined oogonium
- stops at birth
- secondary oocytes are considered haploid already but when fertilized w
sperm cell, it proceed to meiosis II w diploid
- upon fertilization, it will have 46 chromosomes
- upon fertilization via spermatogonia, it will proceed to anaphase
then separating the ootid from the polar body
- only primary oocytes stay in the ovary
- secondary oocytes are in the fallopian tube (ready for fertilization)
- follicles differ in layers (primordial, primary, growing, mature)
- vesicular follicle (cavity is present; releases secondary oocytes)
- corpus luteum (comes after secondary oocytes) -> corpus albicans
(degenerates from corpus luteum)
- yolk is formed during the production of primary oocyte - cytoplasm of
different germ layers
- growing, maturing, mature oocyte – depending on size
- growing oocyte - oogonium surrounded by follicle
- maturing oocyte - developing antrum - filled with follicular fluid
- mature oocyte - large oocyte
- follicle cells prevent primary oocytes from undergoing meiosis
- upon puberty, females produce sex hormones; primary oocytes ->
secondary oocytes
- gonadotropin – hormone that has action on gonads, not produced BY
gonads
Dictyate State
- sort of ‘statis’ where the developing oocyte arrests in diplotene of the 1​st​ meiotic
prophase following initial maturation while the female embryo is in her mother’s
womb
- mediated meiosis stabilizing factor secreted by the follicle cells of the primordial
follicle
- at the beginning of a menstrual cycle a number of oocytes in primordial follicles
are stimulated by pituitary gonadotropins to continue their maturation
- due to their luteinizing hormone concentration
- either blocks/deactivates the meiosis stabilizing factor
- as a result, egg maturation continues and meiosis 1 occurs
- in many species (including humans), the oocyte then arrests at metaphase
of meiosis II until after fertilization

Vitellogenic Phase
- formation of yolk
- estrogen stimulates synthesis of vitellogenins in liver or equivalent organ, and is
then transported to ovary by circulatory system
- follicle cells transfer this into egg (oocyte)
- molecular structure of vitellogenins is modified in the egg
- deposition of yolk in cytoplasm is mediated by enzymes, endoplasmic reticulum,
golgi bodies, and mitochondria
- yolk platelets are formed

EGG CLASSIFICATION
A. by amount of yolk
1. polylecithal/ megalecithal/ macrolecithal
- huge amount of yolk (birds, reptiles, bony fish)
2. mesolecithal
- medium amount of yolk (amphibians)
3. microlecithal/ oligolecithal
- very little yolk (most mammals)
B. by distribution of yolk
1. telolecithal
- yolk distributed in gradient, concentrated toward one pole of egg,
usually the vegetal pole (e.g. amphibians)
2. isolecithal
- yolk evenly distributed throughout egg cytoplasm (e.g. sea urchins,
human)

endoderm – gastrointestinal tract, digestive system


ectoderm – skin, CNS
mesoderm – notochord, internal organs
HORMONES
Reproductive cycle
- prepare both male and female gametes for fertilization

Hormonal control of reproduction


● Hormone
- chemicals that regulate bodily functions
- made of proteins/ lipids
- produced by organs (endocrine gland – secreted in the blood then reaches all
parts of the body except for places with a blood barrier)
- exocrine glands - part of the gland is secreted containing the hormone, sent to the
target organ
- tropism – action, indicating growth or turning movement of a biological
organism

● Vertebrate reproduction
- cyclic activity
- often related to changing seasons
- environmental cues -> hormonal control -> reproductive cycles (reproductive
process)
- FSH (follicle stimulating hormone) and LH (luteinizing hormone) ; target are
testes and ovaries (gonadotropic hormones/ gonadotropins)

Categories
● tropic - specific organ
● non-tropic - work on multiple organs/cells
● both - one growth hormone

cyclic reproductive patterns of mammals


2 types:
1. Estrous cycle
- lower vertebrates
- associated with more pronounced behavioral cycles than menstrual cycles
- copulation can only occur during the actual reproductive phase of the cycle
(window period); cannot occur beyond
● estrus: period of sexual activity
- the only time the condition of the vagina permits mating
Stages of Estrous cycle
- marked by changes in the uterus, ovary, and vagina
1. Proestrus – period of preparation (follicles grow)
2. Estrus – when mating occurs (within the time- range of ovulation) - if successful
- pregnancy proceeds
3. Metestrus – period of repair
4. Diestrus – becomes small and anemic

Frequency of estrous cycle during the breeding season varies

● Monoestrous – single EC in a breeding season


- ex. dogs, foxes
● Polyestrous – recurrence of EC in a breeding season
- ex. mice, rabbits, squirrels
2. menstrual cycle
- anthropoid primates
- refers specifically to the changes that occur in the uterus, preparedness of the
uterus
- also called the uterine cycle
- caused by the cyclic events that occur in the ovaries, the ovarian cycle

The gonadal steroids and their control


1. ovaries – estrogen
2. testes - testosterone
● GnRH - gonadotropin- releasing hormone
- hypothalamus – secretes it
- gonadotropin-releasing hormone
- hypothalamo-pituitary portal vessel
- stimulate anterior pituitary to release GnRH

● Anterior Pituitary
- FSH (blocks meiosis in primary oocytes and spermatocytes) – Sertoli cells –
stimulate spermatogenesis – inhibin
- LH – Leydig cells – produce testosterone – reprod. tract and other organs
(secondary characteristics)
- Testosterone – auto-gonadotropin; produced by the gonads but exert action on
the gonads as well (on Sertoli cells – stimulate spermatogenesis)
- inhibin – negative feedback; inhibit gonadotropins -

hypothalamus – anterior pituitary – testes (NEGATIVE FEEDBACK)


in ovaries (positive feedback)

DIAGRAM (PHASE): top: ovarian, bottom: uterine

endometrium – outermost layer of the uterus


Variations in the length of mc/ phases
- most cycles: 25-30 days
- > 25d <30d
- Younger women > older women
- 15-19 years old = 30 d
- 30 years old = 30 d
- 35 years old = 28 d
- 25,28, 30 day cycles

Summary of Hormone Effects


● FSH
- stimulates growth and development of the follicle
- stimulates secretion of estrogen
- enhances effect of LH in stimulating ovulation
● LH
- stimulates the final development of the follicle
- stimulates ovulation
- stimulates the development of corpus luteum
- stimulates production of progesterone
● Estrogen
- stimulates repair of uterine lining
- at high concentration, inhibits FSH, however during pituitary hormone
surge, it stimulates further FSH production
- as concentration peaks, stimulates release of LH
● Progesterone
- maintains uterine lining
- inhibits release of FSH
- inhibits release of LH
- fall in concentration results in menstruation
- fall in concentration removes inhibition of FSH and a new cycle begins
FERTILIZATION
sexual reproduction – union of sperm and egg cell, can take place outside (lays eggs) or
inside
fertilization
- beginning of a new organism
- external fertilization (ex vivo)
- internal fertilization (in vivo)
- needs copulation/ mating
- amniotic egg – egg that contains their own fluid
- union of male and female gametes
insemination - deposition of sperm cells into the female reproductive tract

Major events in fertilization


a. contact and recognition between sperm and egg
- acrosome contains hyaluronidase – “ase” – enzyme
- hyaluronic acid – glues cells and tissues together
- actin – contractile proteins, what makes up the centrioles during mitosis
or meiosis, behind the acrosome
- nucleus is behind the actin
- tail of sperm – flagellum
- haploid nuclei – pronucleus
- zona pellucida – jelly coat, hyaluronic acid
- plasma membrane, vitelline layer, zoona pellucida (outwards)
- sperm cell will come in contact with egg cell, acrosome will release the
hyaluronidase that contains/digests hyaluronic acid (ACROSOMAL
REACTION) that will dissolve the jelly coat, exposing sperm-binding
proteins, actin will attach vitelline membrane containing the
sperm-binding proteins, pull the sperm toward the egg
- contact -> acrosomal reaction -> contact and fusion of sperm and egg
membranes -> entry of sperm nucleus -> cortical reaction
- cortex granules – located behind the plasma membrane, will open/burst
releasing their contents in the space between the vitelline membrane and
plasma membrane – perivitelline membrane
- fusion of the plasma membrane
- cortical reaction – cortex granules are released, destroy sperm-binding
proteins, harden the jelly coat
- becomes fertilization embryo that prevents from other sperms from
entering

b. regulation of sperm entry into the egg

- polyspermy – prevented by sodium influx


- *remember chemical in ppt*
- phosphor lipase C – breaks down lipids, calcium release (2 effects),
cortical reaction, trigger interphase stage of mitosis
- fusion of 2 plasma membranes -> cortical granules will harden the jelly
coat and will remove the sperm binding proteins
- upon fusion of the 2 plasma membranes, sodium influx
- upon fusion of the 2 plasma membranes, phospholipase C – enzyme that
breaks down lipids -> calcium release (2 effects) -> 1. cortical reaction ->
2. trigger interphase stage of mitosis

c. fusion of the genetic materials of sperm and egg


d. activation of egg metabolism to start development/metabolic activation of the
oocyte

Sea urchin Fertilization


Transport of gametes and fertilization in mammals
- through insemination
- the sperm cell will reach the cervix in the vagina
- pH gradient – diff concentrations, successive intervals – from 5 to 8 (female)
- sluggish motility of sperm except in the fallopian tube
- capacitation of sperm – became hyperactivated in the fallopian tube (rubbed in
cilia)
- oocyte transport in the female reproductive tract
- ampulla of the fallopian tube – site of fertilization

Sperm transport in the female reproductive tract


- site of insemination
- barriers in the reproductive tract
- uterus – uterine contractions are important in the success of the zygote
- entrance into the uterine tube
- final destination
- capacitation – brushing of the sperm cell along the sides of the fallopian tube
Human female reproductive tract illustrating stages of male gamete transport

● A. Sperm entering cervical mucus at external os of cervix


● B. Sperm interacting with endosalpingeal epithelium of the fallopian tube
● C. Hyperactivated motility of sperm in the fallopian tube
● D. Oocyte in the ampulla of the fallopian tube

Union of gametes
- in mammals
- hyaluronidase is released from the acrosomal cap, digests the corona radiata, ZP,
PM
- sperm specific receptor presenter in the ZP
- ZP3 proteins in rodents
- amphimixis – pronuclei fusion
Accomplishments of fertilization
- completion of the second meiotic block
- restores normal diploid number of chromosomes
- sex of the future embryo determined
- variation
- metabolic activation of the egg

​sperm cells carry the sex chromosome! (xy)


CLEAVAGE
fertilization(zygote) -> cleavage
Cleavage mitosis
- no transcription – just divide cytoplasm of maternal ovum, splitting egg cells in
smaller parts until you run out of mRNA and then it will shift to regular mitosis
- “synchronized cell division”
- no G1

Functions of Cleavage Cell Division

1. generate large number of cells


2. generate many copies of the zygotic genome
3. can result to cytoplasmic gradients
- the two-cell would have more cytoplasmic content than the
eight-cell stage
4. increases the nuclear-cytoplasmic volume ratio
- universal aspect of cleavage
- at a certain level, pace of cell division shows
What fuels cleavage?
MPF – mitosis promoting factor (translated from maternal mRNAs)
- fusion of 2 subunits:
● Cyclin B
● cdc2/cdk1 = cyclin dependent kinase (enzyme that allows
transfer of phosphorus/phosphate -> phosphorylation)
- degradation of cyclin in cleavage but not in regular mitosis
How does MPF work?

● Synthesis and degradation of MPF ------> cycling of cells


Activation/deactivation of cd kinases

Cyclin B-cdc2 complex (active MPF)


- phosphorylates target proteins inside the cell
● histone proteins (chromatin condensation)
● nuclear membrane proteins (nuclear membrane depolarization)
● reg. cytoplasmic proteins (spindle fiber organization)

Sources of MPF in the early embryo


- Cyclin B is translated from maternal RNAs
- all the other necessary components for cell cycling are also maternally
derived
- the cell cycle is independent of the nuclear genome for several cell divisions
- regulators of Cyclin B resides in the cytoplasm of the egg

● As the egg cytoplasm and the maternally-loaded cyclin B and other cell cycle
factors are depleted
● Zygote transcription – must begin in order to
replenish them, transcribe their own mRNA
● transition from maternal to zygotic genome
transcription
Beginning of new phenomena:
1. addition of gap phases
2. loss of synchronous division
3. new mRNAs are transcribed
4. new cells synthesize different regulators

When does the embryo stop dividing?


- a new balance in the nuclear:cytoplasmic volume ratio
- depletion in the maternal reserve of mRNAs and proteins
- Result to:
● zygote genome activation (ZGA)
- new gene transcription in the zygote nucleus
● synchronous cell division is lost (becomes asynchronous)

The Timing of Cleavage Division

Cell cycle of a typical mature somatic cell


- includes M (mitosis) phase, S phase (DNA synthesis)
- two gap phases, G1 and G2 - when cells grow, produce mRNAs and proteins
necessary for cell cycle
Cell cycle of early cleavage divisions
- gap phases do not occur
- the cells cycle between M phase and S phase (biphasic)
- if G2 would occur, it is short
- there is not growth
What controls the cell cycle

Cell cycle (cleavage stage embryo) Cell cycle (typical somatic cell)

In normal cell cycles


- presence of the cyclin-cdc2 complex
- promotes the transition from G2 to M phase
In early cleavages
- the cyclin-cdc2 complex promotes the transition from S to M phase

Factors influencing Cleavage Patterns


1. Maternal cytoplasmic factors
angle of mitotic spindle -> site of 1st cleavage furrow
- invariably, cytokinesis occurs in a plane perpendicular to the axis of
mitotic spindle
- maternal gene products may orient mitotic spindle
- prophase occurs because of phosphorylation
-
2. Amount and distribution of yolk
● isolecithal/oligolecithal
● mesolecithal
● telolecithal
● centrolecithal

Patterns of Cleavage and Cleavage Symmetry

Holoblastic radial

Holoblastic rotational

In mammals
- cleavage is extremely slow; during its journey down to the oviduct
- unusually asynchronous
- does not always proceed regularly from 2->4->8 blastomeres
Mammalian Cleavage to Blastula

1. Compaction
- 8 cell stage
- takes place when the cleavage cells/blastoemeres flatten to become more
compact
2. Polarization
- around 16 cell stage, outside cells (9-14), inside cells (2-7)
- concentrated on one side of the blastula/blastocyst - cells separating into
2 poles: embryoblast (inner) and trophoblast (outer)
3. Cavitation
- blastocoel
- holoblastic – blastocyst, whole zygote splits up, usually occur among
animals who develop inside their mother’s womb
- meroblastic – partial, only top portion undergoes cleavage, retains a huge
amount of yolk for nourishment of the growing embryo, takes place
outside the mother’s womb, blastodisc, blastoderm

GASTRULATION
Syngamy – fusion of 2 cells

Gastrulation
- depends if the cleavage is holoblastic or meroblastic
- embryo prepares to segregate into 3 germ layers
- gastrula – gut
Fertilization
- in all sexually-reproducing animals, the first step is fertilization – union of male
and female gametes
- fertilization itself consists of 3 events:
● sperm penetration and membrane
● egg activation
● fusion of nuclei
Sperm penetration and membrane fusion
- protective layers of egg include the jelly layer and vitelline envelope in sea
urchins, and the zona pellucida in mammals
- the acrosome of sperm contains digestive enzymes that enable the sperm to
tunnel its way through to the egg’s cell membrane
- membrane fusion permit sperm nucleus to enter directly into egg’s
cytoplasm
Cleavage
- rapid division of the zygote into a larger and larger number of smaller and
smaller cells called blastomeres
- not accompanied by an increase in the overall size of the embryo
- two embryo ends are:
- in a blastocyst:
● animal pole
- external tissues
- upper portion
● vegetal pole
- internal tissues
- lower portion
- outermost blastomeres in the ball of cells become joined by tight junctions
- innermost blastomeres pump Na+ into the intracellular spaces
- create osmotic gradient, which draws water
- the result is a hollow ball of cells, the blastula, containing a fluid-filled cavity,
blastocoel
Cleavage Patterns
- eggs with large amounts of yolk undergo meroblastic (incomplete) cleavage
- in eggs of reptiles and birds, the clear cytoplasm is concentrated at one
pole called the blastodisc
- cleavage is restricted to this area - resulting embryo is not spherical
- mammalian eggs contain very little yolk and undergo
- from a blastocyst:
● trophoblast
- becomes placenta
- outer layer
● inner cell mass
- forms the developing embryo
- located at one pole
Cleavage Patterns

Fate of Blastomeres
- in mammals, early blastomeres do not appear to be committed to a particular fate
- the earliest patterning events occur at the 8-cell stage
- outer surfaces of the blastomeres flatten against each other in a
process called compaction
- produces polarized blastomeres which then divide
asymmetrically
Gastrulation
- embryo prepares to segregate itself into 3 germ layers (diff organs will arise from)
- gut will be the first to form
- process involving a complex series of cell shape changes and cell movements that
occurs in the blastula
- invagination (cell sheet dents inward), involution (rolling inward), ingression
(breaking away from cell sheet and migrate as individual cells ; for primitive
streak), delamination (cell sheets split in 2)
- establishes the basic body plan and creates the 3 primary germ layers: ecto, meso,
endo

- cells move during gastrulation using a variety of cell shape changes


- cells that are tightly attached to each other via junctions will move as cell sheets
● invagination - cell sheet dents inward
● involution - cell sheet rolls inward
● ingression - cells break away from cell sheet and migrate as individual cells
● delamination - cell sheet splits in 2
Developmental Fates of the Primary Germ Layers in Vertebrates
- epidermis and nervous system – ectoderm
- digestive tract, gut – endoderm
- remaining organs – mesoderm

4 gastrulation patterns
- vary according to amount of yolk
1. sea urchin – mesoderm form first, then endoderm
- follow hollow symmetrical blastulas
- develop from relatively yolk-poor eggs
- deuterostome - anus then mouth
- formation of vegetal plate – ingression of primary mesenchyme cells
(future mesoderm cells) into the blastocoel
- remaining cells invaginate into blastocoel – forming endoderm
- archenteron – future digestive gut
- cells staying at the surface form ectoderm
- involution, ingression, invagination
2. frogs – endoderm forms first, then mesoderm
- from blastula, cells from animal pole involute over dorsal lip of blastopore
into the blastocoel
- cells eventually press against far wall
- eliminate blastocoel – producing archenteron with yolk plug
- movement create layers
- outer ectoderm and inner endoderm
- mesoderm forms later in between
- involution of the animal pole

3. birds – ectoderm forms first, then mesoderm


- undergo meroblastic cleavage
- avian consists of a disc of cells
- instead of blastocyst, blastoderm (on top is blastodisc), sitting on top of a
large yolk mass
- 1st, blastoderm delaminates into 2 layers (endoderm and ectoderm) w/
blastocoel cavity
- upper layer produces 3 germ layers:
- cell that migrate through primitive streak form
endoderm/mesoderm
- cells that remain form ectoderm
- delamination (main cellular movement) and ingression
ingression – from ectoderm
hensen’s node will become the nervous system
inner cell mass of mammals are the blastodisc in birds

4. mammals
- process similar to birds
- embryo develops from inner cell mass
- ICM flattens and delaminates into 2 layers
- a primitive streak forms
- cell movements through it give rise to 3 primary germ layers
- holoblastic

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