Vertebrate Morphogenesis:

General
Definition
- biological process that involves shape development
- Biogenetic Law (Heckel’s law)
• theory of development
• ontogeny recapitulates phylogeny
• development, especially the earlier stages, tell us about evolution

Morphogenic movements
1. Epiboly
Y Overgrowth of 1 CELL LAYER by another
Y Spreading and increase in SA of a sheet of cells to occupy the
entirety of the cell

2. Invagination
Y Formation of a pocket
Y Impocket or infold
Y Sliding or gliding of a mass of cells from the surface to interior, BUT not
over the blastopore lip
Y Involves buckling of cells
3. Delamination
Y Sheet of cells splitting into parallel layers
4. Involution
Y Rolling in of a surface layer over a blastopore lip
Y No buckling of cells

5. Ingression
Y “tearing away” of free cells from a previous layer and later
development into another layer

6. Elongation
Y Lengthening of cells; cytoskeletal involvement

7. Intercalation and Convergence

Y Movement to the midline or center
Y Convergence – narrowing of cells along an axis
Y Intercalation – fusion of cell layers in which cells from each previous
layer alternate and exchange places
Y Convergent extension – elongation in a perpendicular axis
Egg Classification
1. Yolk content
a. Microlecithal – blastula is a hollow ball of cells with a blastocoel; higher
mammals form a blastocyst with a blastocoel, inner cell mass, and
trophoblast; BARELY ANY YOLK
• Amphioxus (no yolk), Placental animals

b. Mesolecithal – blastocoel is displaced into animal pole; YOLK DIVIDES

SLOWLY
• Amphibians, teleosts, lampreys

c. Macrolecithal – Blastoderm forms at the animal pole; YOLK DOES NOT

DIVIDE
• Sharks, Monotremes, Birds and Reptiles

2. Yolk distribution
a. Isolecithal – yolk is equally concentrated throughout
• Urochordates, cephalochordates, and therians
b. Telolecithal – yolk is concentrated in the vegetal pole
• all mesolecithal and macrolecithal vertebrates

3. Egg disposition
a. Oviparous – eggs shed to environment (macrolecithal) or as eggs with
larvae (mesolecithal) and obtain nourishment from yolk
b. Viviparous – eggs retained during development in reproductive tract
i. Ovoviviparous – mother gives oxygen and protection, individual
yolk gives nutrition (SNAKES, SOME SHARKS, SOME FISH, INSECTS)
ii. Euviviparous – obtain constant nourishment from maternal tissue
or placenta
***viviparity evolved in all craniates EXCEPT agnathans, birds, corcodiles
and turtles***

Egg Fertilization
 • Spermatophore (urodeles) – gelatinous packet of sperm deposited
near females, their cloacal lips pick it up (unless deposited directly in
cloaca)
• Spermathecae – sperm inside a tough coat that remains in
reproductive tract, and is only released when female ovulates

Early Embryonic Development

- induced by FERTILIZATION
- Generalized steps:
1. Initiation of cleavage
∆ No increase in cytoplasmic mass and no growth phase (G1 and
G2) between successive divisions (unlike mitosis)
∆ Morula – 16-cell stage and above (until blastula)

2. Blastula (Blastoderm, Blastodisc, or Blastocyst)

∆ Morula cells produce Na+ that causes uptake of water and
induces formation of cavity
∆ Blastocoel

3. Gastrulation (and Neurulation)

∆ Induced by morphogens
∆ Rearrangements and migrations
∆ Formation of 3 germ layers
∆ Establishment of bilateral symmetry
∆ Notochord formation

4. Extraembryonic Membranes
5. Organogenesis

Other Terms
o Morula – roughly 16-32 cells
o Blastodisc – blastula of reptiles and birds; ICM derivative in mammals
o Blastoderm – blastula of amphibians
o Blastocyst – blastula of therians
Trophoblast – outer layer around blastocoel; absorptive; precursor to
chorion and placenta formation + does not contribute to
development of embryo
Inner Cell Mass – at 16-cell stage; forms embryonic or blastodisc
o Gastrocoel or Archenteron – primitive gut formed by migration of
ectoderm, which becomes endoderm
o Neurula – product of neurulation; usually occurs before a complete
gastrula is formed
o Neurocoel – cavity of neural tubes
o Epimere – upper layer of blastodisc/blastoderm blastomeres; full
participation in embryogenesis
o Hypomere – lower layer of blastodisck/blastoderm blastomeres; doesn’t
participate in embryo formation

I. CLEAVAGE
- 1 cleavage is ALL THE SAME (Perpendicular to each other; meridional)
st

- 2nd cleavage differs (different forms of equatorial cleavage)

» Sometimes asynchronous (makes odd number of cells)

» determined by yolk distribution
a. Holoblastic– complete division
¬ Isolecithal è Equal sizes of blastomeres
Radial – echinoderms, Cephalochordates
Spiral – Annelids
Bilateral - Ascidians; Urochordates
Rotational – mammals, nematodes
§ 1st cleavage = meridional à 2 cells
§ ½ cells divides meridionally
§ ½ cells divides equatorially
¬ Mesolecithal è unequal sizes of blastomeres
Radial – amphibians
b. Meroblastic (Macrolecithal)– incomplete division; involves
Micromeres (smaller, animal) and macromeres (larger, yolk)
Ω Formation of the Subgerminal Space (fluid-filled cavity
between the blastoderm and yolk)
Ω Vegetal pole cells DO NO DIVIDE

Discoidal – very small micromeres at animal pole, and

very large yolk mass of macromeres
∞ Birds, Reptiles, monotremes, fish

Superficial – mitosis but not cytokinesis resulting in

multinucleated cells
∞ Insects
II. BLASTULA
 III. GASTRULATION
» “Gut formation”
» Differs from cleavage because there is REARRANGEMENT
» Main events:
√ Archenteron formation (primitive gut)
√ Bilateral symmetry
√ Neurulation occurs simultaneously
√ 3 germ layers formed
√ extraembryonic membranes
» Coelom is formed as the space between mesoderm layers
» Varies per species

Ectoderm
 - ALWAYS the first tissue formed
- Precursor of other 2

Endoderm
- begins as endomesoderm
- encloses the archenteron
• grows downward to enclose gastrocoel DURING notochord
formation

- future outpouchings or diverticula become other organ linings

- EPITHELIAL in nature
Mesoderm
- Begins as endomesoderm
- do not form a sheet
- produce loosely connected cell clumps à MESENCHYME (totipotent)
- Mesodermal regions with individual cavities
• Notochord (chordamesoderm)
• Lateral mesoderm sheets (lateral to noto) – differentiates into:
a. Epimere (paraxial mesoderm) – myocoel
Ω Somitomeres – loosely held mesenchyme from the neck to
posterior end; form the somites when they start detaching
à body wall musculature
Ω Cephalic somitomeres remain connected à muscles of
face, jaws and throat
Ω Gives rise to: Cardiovascular system and outer lining of
organs
b. Mesomere (intermediate/nephrogenic mesoderm) – nephrocoel
Ω Gives rise to: Kidneys
c. Hypomere (lateral plate mesoderm) – coelom
v Mesoderm grows downward and delaminates
Ω inner wall = splanchic mesoderm (+ endoderm =
splanchopleure)
Ω outer wall or body wall = somatic mesoderm (+ ectoderm =
somatopleure)
Ω gives rise to: striated muscles and dermis, gonads
- Cavity formation
Ø Enterocoely – primitive; mesoderm pinches off and has a cavity
(cephalochordates and lampreys)
Ø Schizocoely – mesoderm forms as a sheet, then splits (everything else)
IV. NEURULATION
» Aided by expression of HOX gene
» Results in Neurula
» Primary neurulation (tetrapods)
▽ Ectoderm thickens into neural plate à neural folds at the
sides à move up and meet to form a neural tube
∞ Schizocoelous
▽ Before neural fold fusion, some cells pinch off as Neural crest
cells
Head à sheets
Trunk à cords
▽ Neural tube differentiates into Brain and SC
» Secondary neurulation (lampreys)
▽ Ectoderm thickens into neural plate à sinks into midline
dorsal surface as a rod à neural keel à cavitation à neural
tube
∞ Primitive; enterocoelous
▽ No neural crest cells from neural folds
▽ Neural crest cells – pinch off from lateral parts of neural keel

V. EXTRAEMBRYONIC MEMBRANES
1. Amnion – innermost sac enclosing the embryo; outgrowth of
somatopleure; for cushioning (NONE IN FISH AND AMPHIBIANS)
2. Yolk sac – Simple yolky placenta in ventral region for nutrition; highly
vascularized; secreted yolk is brought to embryo via vitelline veins;
SHRINKS AS EMBRYO GROWS; Vestigial in mammals
3. Chorion – eggshell lining of uterus; outgrowth of somatopleure; gives
rise to placenta and made by the trophoblast; for oxygen access
(ovi) or nutrient access (vivi)
4. Allantois – midventral evagination of cloaca; uric acid storage that
fuses with chorion (chorioallantoic membrane) near eggshell inner
surface for better access to gas (BIRDS, MONOTREMES, AND REPTILES)
or fuses with placenta (chorioallantoic placenta) for gas, nutrients,
and waste segregation (MAMMALS); gives rise to umbilical cord and
part of GI (MAMMALS)
∆ Part near cloaca è bladder
∆ Part far from cloaca è urachus
∆ Forms diverticulum in ventral hindgut
5. Placenta – maternal tissue + embryonic membrane; connected via
umbilical cord
§ Chorioallantoic placenta (eutherians) – deciduous
(membranes peel endometrium at birth causing bleeding) or
non-decidious (fetal embryonic membranes peel from
endometrium without bleeding or shedding)
 bird reptile

VI. ORGANOGENESIS
Specialized cells
1. Neural Crest (ectomesenchyme) – somatic and autonomic ganglia, facial
bones; ectodermal origin
2. Epidermal or Ectodermal placodes – sensory organs\
Ectoderm sinks beneath skin and gives rise to neuroblasts and
sensory epithelia
 Nasal placodes, otic placodes, optic placodes (lens), epibranchial
placodes (Cranial nerves), Linear placodes

Morphogens
• dorsal lip of blastopore à organizer area: establishes longitudinal
axis
• Proteins that induce differentiation
• Homeotic genes – segmental genes that determine fates of germ
layers
Y Example: HOX GENE CLUSTERS = Segmentation of head
Y Example: Hedgehog gene = notochord differentiation
• Chordin – axis-forming activity during gastrulation; encoded by
dorsal lip and ONLY encoded in the tail region
• Induces mesenchyme differentiation
∆ Mesenchyme à aggregates into blastemas à give rise to
general organ architecture

Vertebrate Morphogenesis:
Variations

AMPHIOXUS
- Microlecithal; isolecithal; holoblastic radial

Cleavage and Blastulation

1st cleavage à 2nd cleavage perpendicular à 3rd cleavage semi-equatorial

(just above equator) and perpendicular to the first 2 à 8-celled morula

asynchronous division of morula à 32-celled blastula with blastocoel

Gastrulation and Neurulation

Invagination of vegetal wall (presumptive endoderm) à inner cells of

invagination form the endomesoderm à moves further towards opposite side
ectoderm à blastocoel disappears, gastrocoel and blastopore formed

parts of Presumptive mesoderm pinch off upward as hollow balls (enterocoely)

à mesodermal pouches or coelomic pouches à their cavities meet to form the
coelom
***at this time, it is hatched as a free-swimming larva***

Dorsal midline mesoderm differentiates à chordamesoderm à induces

notochord formation and CNS formationà dorsal ectoderm thickens into
neural plate à edges meet and form a hollow neural tube
Mesodermal bands grow ventrally à establish somatopleure and
splanchopleure à differentiates into epimere, mesomere, and hypomere

FISHES (LAMPREYS AND TELEOSTS)

- Macrolecithal; Telolecithal; Meroblastic discoidal

Cleavage and Blastulation

More divisions at animal pole à vegetal blastomeres large and yolky, animal
blastomeres smaller and more numerous à Animal blastomeres form
blastoderm à small blastocoel between blastoderm and vegetal pole
Gastrulation and Neurulation

Indentation at dorsal region à dorsal lip of blastopore à epiboly then involution

à entering cells become endomesoderm à gastrocoel forms and blastocoel
disappears

Blastoderm grows over the yolk surface by epiboly à become yolk sac
enclosing vegetal cells

Posterior blastoderm concentrates à embryonic shield à becomes embryo

later on

Endomesoderm à endoderm + mesoderm

Mesoderm à forms chordamesoderm at midline à become notochord and

lateral mesodermal plates growing around yolk

AMPHIBIANS (REP: FROG)

Cleavage and Blastulation

More divisions at animal pole à vegetal blastomeres large and yolky, animal
blastomeres smaller and more numerous à Animal blastomeres form
blastoderm à small blastocoel between blastoderm and vegetal pole

Gastrulation and Neurulation

Epiboly à blastopore à involution of cells over the blastopore lip à

endomesoderm and yolk plug
Middorsal Endomesoderm à chordamesoderm à LATER notochord

Rest of the endomesoderm à paired lateral projections grow upward à meet

just below notochord à endoderm

other parts of endomesoderm à mesoderm proper à grows downward

between ecto and endoderm à schozocoely à somato and splanchopleure

ectodermal thickening à neural plate à extends upward à hollow neural tube

mesoderm à epimere, mesomere, hypomere

REPTILES AND BIRDS
- discoidal
- No blastopore; primitive streak functions as a blastopore

Cleavage and Blastulation

Animal blastomeres à blastoderm (blastodisc) à epiblast and hypoblast with

subgerminal space

Gastrulation and Neurulation

Epiblast à thickens at posterior à Hensen’s node in caudal end (primitive

node) leading to a primitive streak

Epiblast à epiboly to primitive streak à involution and ingression à (a) migrate

to the subgerminal space (b) migrate to surface of hypoblast à (a) mesoderm
(b) endoderm

hypoblast cells displaced to peripheries à become endodermal yolk lining

Epiblast cells that did not migrate à ectoderm

Hensen’s node à start of notochordal process growth à Mesoderm grows

alongside it à lateral to notochord à epimere, mesomere, hypomere

***mesoderm grows except in cranial region; ectodermal***

Ectodermal thickening à neural plate à fold up à neural tube

Hypomere à splits à somatic and splanchic mesoderm à schizocoely à

coelom

***same in reptiles, but blastopore is retained!!!

MAMMALS (MONOTREMES)
*same as birds*

- modification: embryo becomes bigger due to uptake of uterine nutrients

BEFORE being enclosed in an egg shell such as with birds
MAMMALS (MARSUPIALS)

Cleavage and Blastulation

Blastodisc – DOES NOT have ICM or does not form a blastodisc

Unilaminar protoderm à spread around layer of blastocyst à have a

trophoblast

Gastrulation and Neurulation

Protoderm à bilaminar à ecto and endoderm à primitive streak forms on

ectoderm à usual involution and ingression to form mesoderm

MAMMALS (EUTHERIANS)
- Microlecithal, holoblastic, rotational

Cleavage and Blastulation

Cleavage within Zona pellucida and follicles à morula à blastocoel + ICM +

trophoblast à blastocyst

Arrival in uterus à Zona pellucida secretes blastocyst à trophoblast attaches

ICM à embryonic disc è epiblast and hypoblast

Gastrulation and neurulation

*same as bird, but from the ICM*