Trans - Patten's Foundations of Embryology Chapters 1-7
Trans - Patten's Foundations of Embryology Chapters 1-7
Trans - Patten's Foundations of Embryology Chapters 1-7
Special Fields
Historical Background
Galen
- learned about structure of relatively
advanced fetuses
- minute dimensions resisted serious
analysis
- development of microscope allowed
study of early stages
de Graaf
- described ovarian follicles
Comparative Embryology
- Provided insight for concept that
ontogeny recapitulates phylogeny
- Recognition of different modes of
development
- Adoption of model organisms
Descriptive Embryology
- Basic structural
embryonic body
- Serial
sections,
reconstructions
pattern
3D
wax
of
the
plate
Experimental Embryology
- Understand
causative
factors
in
development by posing hypotheses
and testing them by manipulation of
embryos
- Roux
o Experiment of 2 cell embryo
Destroyed
one
blastomere
Each cell is capable of
giving rise to complete
individual
Provided
proof
of
untenability
of
the
preformationist doctrine
Chemical Embryology
- Provided description about chemical
and physiological events in embryo
- Interaction between components and
how basic body pattern is laid down
Teratology
- Concerned
with
study
of
malformations
- Identify and eliminate genetic and
environmental factors that cause
congenital defects
Reproductive Biology
- Related to problems of conception and
contraception
- Emphasis
on
gametogenesis,
endocrinology, transport of gametes,
fertilization, embryonic development
Developmental Biology
- Embryonic development + postnatal
development and processes
- Focus on processes and concepts,
rather than specific morphological
structures
- Plant and animal systems included
Embryology in Contemporary Society
Test tube baby
In vitro
transfer
fertilization
and
embryo
Golgi
complex
membrane vesicles
- May
be
mediated
by
receptor
molecules located at cell surface
activated by binding of a ligand,
causing
stimulation
of
signal
transduction.
Cell Surface
- Junctional complexes
o Desmosomes
bind epithelial cells
focal
points
for
attachment of fibrillar
intracellular proteins
o Gap junctions
Mediates communication
and exchange of small
molecules
o Tight junctions
On surface of many
epithelia
Bind
adjacent
cells
together,
forming
impermeable barrier
Prevent
mingling
of
membrane proteins on
either side of junction
- Cell adhesion
o Experiment: disaggregated and
reaggregation of sponges
o Ca++ mediated adhesion
Glycoproteins
E-cadherin
(epithelium)
N-cadherin
(nerves,
mesoderm)
P-cadherin
(placenta)
o Heterophilic binding between
complementary saccharides
Occurs
during
mammalian fertilization,
when
head
of
spermatozoa encounters
zona pellucida
Extracellular Matrix
- Cells embedded on extracellular
matrix
- Collagen (glycoprotein with glycine)
o Basic unit is tropocollagen
o (I, II, III, IV, V, X)
Differentiation
- Restriction and determination signify
progressive
limitation
of
developmental capacities
- Differentiation: actual morphological
or functional expression of a particular
cell or group of cells; the process
where cell is specialized
Morphogenesis
- Processes that mold external and
internal configuration of embryo
- Pattern formation: laying down of
morphological blueprint
- Morphogenesis: actual realization of
plans
- Homeotic
genes:
determine
the
regional
characteristics
of
each
segment (14 segments)
Gene Activation
- Derepression of heterochromatin or
repression of euchromatin
- Derepression of general genes from
zygote to blastula
- Derepression of tissue specific genes
from gastrula to organogenesis
Restriction and Determination
- Restriction:
the
reduction
of
developmental options permitted to a
cell
- At
gastrulation,
one
stage
of
restriction has occurred (endo, meso,
ectoderm)
- Part of ectoderm thickens and
undergoes Neurulation
- Determination
o restriction has proceeded to a
point where a group of cells
becomes committed to single
developmental fate
o final
step
in
process
of
restriction
- Inductions
(tissue
interactions)
precede
determination
(and
restriction)
Induction
- Form of embryonic signal calling
- Effect of one embryonic tissue on
another, so that the developmental
course of responding tissue is changed
from what it would have been in
absence of inductor
- First major inductive event: induction
of mesoderm in cleavage
- Primary
induction:
induction
of
nervous system during and shortly
after gastrulation
- Secondary induction: nervous system
induces other structures
- Permissive
induction:
inductive
signal
required
to bring about
development of structure
- Instructive induction: responding
tissue has options of forming more
Intercellular Communication
- Intercellular
communication
place in localized gap junctions
takes
Cell Movements
- Individual cells commonly migrate by
means of amoeboid movements
- Unique: in avian embryos, primordial
germ cells move from wall of yolk sac
into the bloodstream and are carried
via blood to the gonads
- Amoeboid movement examples:
o Ectoderm: migration of cells
away from neural crest
o Mesoderm: spreading out of
mesodermal cells
o Endoderm: migration of primary
germ cells from yolk sac to
gonads in mammals
- Movement as sheet seen in epithelial
cells
Cell Death (Apoptosis)
- Examples:
o Tail and opercular resorption
o Separation of digits
- May be hormonally controlled
o Male and female genital ducts
The Clonal Mode of Development
- Clones: group of cells arising from
single precursor
Regulation and Regeneration
- Regulation
o restoration of missing material
occurring before differentiation
of structure
o basis
for
development
of
identical twins
o subdivision of inner cell mass
- Regeneration
o Differentiation
of
structures
already occurred
Growth
- Differential growth: all parts of the
embryo do not grow at the same rate
- Determinate growth: body grows to
certain point that is characteristic of
species and sex (mammals)
- Indeterminate
growth:
ancestral
vertebrates (fish)
Recapitulation
- Biogenetic law of Muller and Haeckel
o Ontogeny is an abbreviated
recapitulation of phylogeny
REPRODUCTIVE
SEXUAL CYCLE
ORGANS
AND
THE
Reproductive Organs
Female Reproductive Organs
- Paired gonads (ovaries) located in
pelvic cavity
- Each ovary lies close to funnel-like
opening (ostium tubae) at end of a
uterine (fallopian) tube
- Around abdominal orifice of tube is
fimbriae
- Uterus:
thick,
vascular,
smooth
muscle, caudally continuous with the
cervix, will project into vagina
- Vagina: organ of copulation and birth
canal
Male Reproductive Organs
- Testes suspended in scrotum (with a
countercurrent
heat-exchange
system), with lower temperature
- Spermatozoa produced in seminiferous
tubules, then to tubuli recti, rete
testis, ductuli efferentes, epididymis,
ductus deferens
- Spermatozoa stored in epididymis and
ductus deferens
Sexual Cycle in Mammals
Estrous Cycle in Mammals
- Sexual cycle
o Estrus
(prepared
for
reproduction accompanied by
ovulation and sexual desire)
o Postestrum
(regression
of
preparations)
o Diestrum (period of rest)
o Proestrum (period of active
preparatory changes)
- Light is a critical factor: higher than
certain threshold causes hypophysis to
become active and produce enough
FSH
- Light Nervous transmission FSH
Follicular growth Estrogen in
ovarian follicle estrogen in blood
stream mating behavior, estrous
uterus, estrous vagina
Primate Menstrual Cycle
- Sexual cycle is characterized by
menstruation
- Commences
at
menarche
until
menopause
- Three phases
o Menses
o Proliferative (follicular)
o Secretory (luteal)
- Menstruation initiated by reduction of
blood flow into superficial uterine
blood
vessels,
resulting
in
deterioration and extravasation of
blood into tissue, bringing with it the
necrotic superficial tissue
stimulation
of
GnRH
and
gonadotropins
Hormonal Regulation of Reproduction in the
Male
- Testosterone: secreted by Leydig cells
(stimulated by LH)
- Sertoli cells take up FSH synthesis
of ABP
GAMETOGENESIS
Gametogenesis
- Germ plasm gametes + cells that
give rise to them
- Gametogenesis:
germ
plasm
is
converted to specialized sex cells
capable of uniting at fertilization and
producing a new being
- Four phases
o Origin of germ cells and
migration to gonads
o Multiplication of germ cells in
the gonads (mitosis)
o Reduction
of
number
of
chromosomes (meiosis)
o Maturation and differentiation of
gametes
The Origin of Primordial Germ Cells and Their
Migration to the Gonads
- Primordial germ cells of mammals,
reptiles, birds arise in epiblast take
up
temporary
residence
in
extraembryonic
tissue
before
returning
o Birds: in germinal crescent
o Mammals: posterior wall of yolk
sac (near allantois)
o Amphibians:
vegetal
pole
cytoplasm
- Note: PGCs do not produce gonads,
they produce gametes
- PGCs in extragonadal sites may
develop into teratomas
Proliferation of Germ Cells by Mitosis
- Mitotically active germ cells: oogonia,
spermatogonia
- Settling in the gonads induce a
proliferative phase
- Mammalian
oogonia
proliferative
phase
o Mitosis brings numbers to about
7 million at 5th month of
pregnancy
o Atresia causes sharp decline
o Primary
oocytes
formed
(suspended at prophase I)
Meiosis
- Genetic recombination occurs by
o Random
distribution
of
chromosomes to daughter cells
o Crossing over
- (2n, 4c) (Me I) (1n, 2c) (Me II)
(1n, 1c)
Spermatogenesis and Oogenesis Compared
- 4 functional spermatozoa vs 1 viable
ovum
- Arrests in meiosis
o Spermatogenesis: none
- Oogonesis meiotic arrest
o First
arrest
at
diplotene
prophase I in primary oocyte
o Broken by hormonal changes
o
o
Spermatogenesis
- Transition from mitotically active PGCs
to mature spermatozoa
- Three phases
o Mitotic multiplication
(spermatocytogenesis)
o Meiosis (spermatidogenesis)
o Maturation and differentiation
(spermiogenesis)
- Spermatogonia
o Type A: stem cell population
Ad: long term reserves
Ap: mitotically active,
give rise to B
o Type B: committed to finish
spermatogenesis
- Spermatocytes
o Preleptotene spermatocytes
o Primary spermatocytes (Me I)
secondary spermatocytes (Me
II) haploid spermatids
- Differentiation
of
spermatid
to
spermatozoa
o Golgi
complex
forms
proacrosomal
granules, into
acrosome
o Centrioles as point of anchorage
for developing flagellum
- Intercellular
bridges:
facilitate
synchronous
differentiation
and
division of sperm-producing cells
- Sertoli cells:
o FSH target Sertoli cells
o Synthesis of ABP (to maintain
high testosterone levels)
o Maintain blood-testis barrier
o Create
environment
for
differentiation of sperm cells
o Facilitate release of mature
spermatozoa
o Degradation of residual bodies
- Blood-testis barrier (held by tight
junctions) responsible for preventing
bodys
immune
system
from
destroying
mature
sperm
cells
(antigenically different from body)
Gene Expression during Spermatogenesis
- Posttranscriptional control
Sperm Maturation
- Sperm coated with glycoprotein which
must
be
removed
in
female
reproductive tract before fertilization
can occur (activation, capacitation)
- Seminal fluid provides external energy
source (causes nonmotile sperm to
become motile)
Oogenesis
Oogenesis in Amphibians
- Mitotic phase of oogenesis does not
come to early halt
- New crop of eggs each year, 3 years
for maturation
estrogen
secretion from oocyte
into liver secretion of
vitellogenin from liver to
oocyte
o Yolk precursor vitellogenin
(phospholipoprotein)
Incorporated into oocyte
by micropinocytosis
Represented
by
phosvitin
and
lipovitellin (packed in
crystalline form to form
yolk platelets)
o Yolk formation was thought to
be function of Balbiani body
(yolk nucleus)
o Pigment granules concentrate
at animal hemisphere
- Maturation phase
o Hormonally induced release of
egg from first meiotic block
o Breakdown of germinal vesicle
o Completion of first meiotic
division
o Formation of first polar body
o Begins
with
secretion
of
progesterone (stimulated by
gonadotropin)
Causes breakdown
of
germinal vesicle, meiotic
maturation
Meiosis
arrested
again
at
Metaphase II by CSF (cytostatic
factor)
Oogenesis in Birds
- Yolk is a single cell (the ovum)
o Gradually
accumulates
in
cytoplasm of ovum before it is
liberated from ovary
o All other noncellular secretions
(egg white, shell membrane,
shell) are contributed as ovum
passes reproductive tract
- Yolk still produced by the liver and
transported via blood to the follicular
cells
surrounding
ovum
(as
in
amphibians)
o 50% water, 33% fat, 16%
protein, 1% carb
o Water NaCl, Ca salts (bone
formation)
o Proteins lipovitellin (binds w/
lipids), phosvitin (binds w/
phosphorus)
- Protuberance containing ovum is
ovarian follicle
- With zona radiata (irregular striated
plasma membrane due to microvilli),
for increase in membrane surface,
enhancing metabolic interchanges
- Compared with mammalian ovum:
o No yolk in mammals, just liquor
folliculi
o Both have two layered CT theca
- Yolk release albumen secretion
shell membranes (isthmus) shell
(uterus)
Oogenesis in Mammals
- Primary oocyte (so called as it is
undergoing meiosis I) + flat follicular
cells = primordial follicle
- Phase I: pool of primordial follicles
developing into primary follicles (flat
cuboidal)
- Meiotic
arrest
follows
(diplotene
meiosis I)
- Both oocytes and follicular (granulosa
cells) develop microvilli, connected by
gap
junctions
(allow
high
MW
molecules to pass through)
- Zona pellucida beings to develop
- Phase II: growth of oocyte and
granulosa covering (under influence of
gonadotropic hormones)
- Overall growth of follicular covering
mediated by FSH receptors
- Secondary follicle when antrum is
identifiable
- LH
receptors
develop,
allowing
production of testosterone by theca
o Transported into granulosa cells,
wherein
granulosa
converts
testosterone to estrogen by
aromatase
Phase III: further follicular growth and
selection of one follicle (highest
receptivity to FSH) which will undergo
ovulation
o Begins late in follicular phase of
menstrual cycle
o After LH surge, angiogenesis
occurs, causing estradiol to spill
out into blood
o Before
ovulation,
ovum
is
released from first meiotic block
(diplotene), allowing meiosis I to
finish
o After that, follicle is now ready
to respond to preovulatory FSH
and LH surge, and release itself
(wherein it is now at metaphase
II, second block)
Ovulation
o Increased antral fluid pressure
within follicle causes bursting of
follicular wall
o Weakening of follicular wall by
lytic enzyme (stimulated by LH)
Corpus luteum
o Stratum granulosum and theca
interna involved in corpus
luteum formation
o Endocrine
organ,
secreting
progesterone and estrogen
o Granulosa
cells
swell
and
develop to secrete high levels of
hormones
o Formation of corpus lutem
require continuous presence of
LH from pituitary
regression happens with
decreased sensitivity to
LH receptors
corpus
lutem
of
pregnancy maintained by
chorionic
gonadotropin
(secreted by embryo)
o Corpus lutem produces large
amounts of progesterone and
estrogen
Progesterone
prepares
lining
of
uterus
for
implantation
Sperm
Penetration
of
the
Egg
in
Invertebrates and the Acrosome Reaction
- When spermatozoa encounter egg,
former undergoes changes
- In presence of egg cells, spermatozoa
will cluster and increase motility
- Direct contact with jelly coat increases
motility and stimulates acrosome
reaction
- Speract is responsible for increased
motility and activated respiration that
occur when sperm contacts with jelly
coat
o Increase in permeability of
plasma causes influx of Na and
Ca, efflux of H+
Raises intracellular pH,
stimulating
flagellar
activity
- Contact w/ jelly coat stimulates
acrosome reaction
o Begins with breakdown and
subsequent fusion of outer
acrosomal
membrane
and
plasma membrane
o Polymerization of G-actin to Factin
(forming
acrosomal
process)
Tip of process is covered
with bindin, mediating
sperm binding to surface
of eggs
o Spermatozoa digest through
vitelline membrane by lysins
Binding of Sperm to the Egg
- Sperm receptor molecule (on microvilli
of egg)
o Intracellular: remains constant
among species
o Extracellular: differs accdg to
species
- After sperm-egg fusion, fertilization
cone forms by microvilli engulfing
sperm head
Blocks to Polyspermy
- Fertilization of egg by more than one
sperm
- Fast block
FERTILIZATION
- Initial contact between egg and sperm
- Entry of sperm cell into egg
- Prevention of polyspermy by egg
- Metabolic activation of egg
- Completion of meiosis by egg
- Formation and fusion of male and
female pronuclei
Mammalian Fertilization
Sperm Transport in the Female Reproductive
Tract of Mammals
- Barriers to fertilization:
o Natural
acidity
of
vagina
(bacteriostatic)
Seminal fluid acts as
buffer (raises vaginal pH)
Seminal fluid may cause
contractions
in
upper
vagina, helping propel
spermatozoa
Orgasms cause uterine
contractions
o Entrance
to
uterine
tubes
(ovulation can sometimes only
occur on one tube)
- Positive rheotactic response (face
an oncoming current generated by
uterine ciliary movement)
- Capacitation:
removal
of
glycoprotein covering spermatozoa,
enabling better penetration of egg
Egg Transport
- Ciliary currents and smooth muscle
contractions transport egg into uterine
tube
- Corona radiata adds mass faster
movement down tube
Union of Gametes
- Mammals: occurs in upper part of
uterine tubes
- Spermatozoa must penetrate corona
radiata cells and then zona pellucida
before contact with plasma membrane
- Zona pellucida: molecules on sperm
head bind with species-specific sperm
receptors (consist of exposed part of
ZP-3)
o Further contact with other core
regions
stimulate
acrosome
reaction
(capacitation
is
prerequisite,
so
that
lytic
enzymes
are
released
to
facilitate passage of sperm
through zona pellucida)
- Acrosome reaction:
o Localized fusion and breakdown
of outer acrosomal membrane
and plasma membrane
o Acrosin
bound
to
inner
acrosomal membrane digests
through zona pellucida
o Fertilization cone bulged out
when sperm makes contact with
egg
Development and Fusion of Pronuclei
- With sperm entry, block to second
meiotic division is lifted, and second
polar body is released
- Breakdown
of
sperm
nuclear
membrane and decondensation of
body
(determines
dorsoventral axis of
future embryo)
Determination of two axes determines
third
Even before cleavage, the three axes
are
established
and
secondary
polarization is completed
Amphibians
- Meridional (begins at animal pole,
bisects gray crescent) meridional
equatorial double meridional
double equatorial
- Amphibian blastocoel
o Formed from specialization of
cleavage furrow of animal
hemisphere
o Filled with Na+ ions, then water
enters
to
maintain
ionic
balance, causing expansion
- Amphibian blastula has three main
regions
o Region around animal pole,
including cells forming roof of
blastocoel
Future ectodermal germ
layer
o Region around vegetal pole,
including large cells in interior
Future endodermal cells
o Marginal ring of cells in the
subequatorial region of the
embryo, including gray crescent
Embryonic mesoderm
- Nieuwkoop
discovered
basis
for
mesodermal induction
o Experiment
1:
directly
apposed sheet of cells from
animal
hemisphere
above
blastocoel to a yolk mass from
vegetal hemisphere
Inductive
influence
from yolk mass caused
animal pole cells to form
mesodermal
structures
Therefore,
blastocoel
may
function to restrict
interaction
between
future
endodermal
and
ectodermal cells
o Experiment 2: isolated pieces
of ectoderm induced to form
mesoderm by transforming
growth factor-beta
Source of mesodermal induction
(Nieuwkoop center or dorsalizing
center
or
mesodermal
inducing
center), resides in a number of vegetal
endodermal
cells
located
in
prospective dorsal midline
Nieuwkoop center
o Stimulates
formation
of
mesoderm
o Establishes dorsal properties of
induced mesoderm
Dorsal induced mesoderm is direct
forerunner of what is called the
Spemann organizer (dorsal lip of
blastopore), the dominant organizing
region of amphibian embryo during
gastrulation
Birds
- Mitotic spindles align themselves so
subsequent cleavage furrows form at
right angles to the preceding one (first
3)
- Fourth
cleavage
furrow
is
a
circumferential one
- Blastomeres formed by first few
divisions are dorsally bound by plasma
membrane but basal surfaces are
open to underlying yolk
- Further cleavage of blastoderm
results in radial extension of embryo
- 32-cell
embryo
shows
cleavage
parallel
to
surface,
establishing
several strata of superficial cells
- At around 100 cells, blastoderm is
underlain by a subgerminal cavity
o pH of subgerminal cavity 6.5,
while albumen: 9.5, leading to
establishment of transepithelial
potential of 25mV. Electrical
gradient
determines
dorsoventral
axis
of
blastoderm.
- Shedding of cells begins from
undersurface of area of blastoderm
that is farthest away from source of
gravity (this area becomes caudal
end of embryo)
o Area pellucida: central portion
of blastoderm thinned out by
shedding of cells
o Area opaca: region where
blastoderm cells still abut
directly onto yolk
- Primary hypoblast: aggregates of
cells shed from lower surface of
blastoderm
by
a
process
of
polyingression/delamination
Mammals
- Equal holoblastic cleavage of an
isolecithal egg
- Persistence of traits characteristic to
large-yolked
embryos
(later
in
development)
- Second cleavage division may not
occur
simultaneously
in
both
blastomeres
o Mitotic
spindle
of
one
blastomere may rotate 90
degrees, causing a rotational
pattern of cleavage
(as
opposed
to
radial
in
echinoderms, Amphioxus)
- 16-cell stage still is contained in zona
pellucida
- Morula: internal secretion of fluid by
blastomeres leads to formation of
blastocoel, or blastocyst cavity
o Similar to amphibian (Na-KATPase system brings in Na, and
then water)
of
Evagination
Involution
Epiboly
Ingression
Polyingression
(delamination)
Ameboid motion
Description
Inpocketing
sheet of cells
Example
of
Outpocketing of
sheet of cells
Rolling around a
corner
of
an
expanding outer
layer of cells and
spreading over an
internal surface
Spreading of a
cell sheet
Sinking
of
individual
cells
from a surface
into an area
Separation
of
second
sheet
from an original
single sheet
Migration of cells
as
single
individuals
through their own
motility
Archenteron
formation
in
Amphioxus
Exogastrulation
Cell movements
through
the
amphibian
blastopore
Spreading
of
outer
cells
towards
amphibian
blastopore
Primary
mesenchyme
formation in sea
urchin embryos
Formation
of
primary
hypoblast
of
avian embryos
Migration
of
neural crest cells
Hensens
node:
notochord
Cranial part of primitive
streak:
embryonic
mesoderm
Caudalmost
part
of
streak: extraembryonic
mesoderm
(trophectoderm)
to
form
endodermal layer of parietal
yolk sac
o Parietal endoderm cells create
basement membrane called
Reicherts membrane
o Polar
trophectoderm:
overlying ICM
can
undergo
mitosis,
daughter cells become
mural
o Mural
trophectoderm:
surrounding blastocyst cavity
Mitosis
results
in
polyploidy giant cells
ICM
undergoes
transformation
different from other mammals
o Protrudes deeply into blastocyst
cavity (like a tongue-like lobe)
o Cavity forms within the lobe
(proamnion),
and
cells
surrounding it are primitive
ectoderm (or epiblast)
Called an inverted egg
cylinder
Neurulation in Amphibians
Formation of the Neural Tube
The Neural Crest