Model organisms in development

Cell and embryology

A few have been studied extensively; each has advantages and

disadvantages.
Model Systems

Xenopus laevis: development is independent (in vitro), easy catch

and observation but poor genetics.
Model organisms: vertebrates (frog, mouse, zebrafish) Chick: available, surgical manipulation and in vitro culture but poor
Model organisms: invertebrates (sea urchin, Drosophila, nematode) genetics.
Mouse: surgical manipulation, good genetics, transgenic model,
Identifying development genes mammalian but development
p is in utero .
Drosophila: great genetics, great development (recent Nobel Prize to
Textbook: Wolpert L, Beddington R, Jessell T, Lawrence P, Meyerowitz E, Smith
J. (2007) Principles of Development. 3th ed. London: Oxford university press. Lewis, Nusslein-Volhard & Wiechaus).
C. elegans: has less than 1000 cells and is transparent.
Gilbert SF. (2003) Development Biology. 7th ed. Sunderland: Sinaure
Sea Urchin : in vitro
Associates Inc.
1 Arabidopsis thaliana: flowering plant. 2

Summary of the main patterns of cleavage

Lecithal

3 4

1
 Model organisms: vertebrates

All vertebrate embryos undergo a similar pattern of development.

1) fertilization
2) Cleavage (cell number ↑, but total mass X)
Fi 2 1
Fig.2.1
3) blastulation (blastcoel formation and three germ layers)
4) gastrulation (where ectoderm covers embryo, endoderm and The skeleton of a mouse
embryo illustrates the
mesoderm are inside), A-P axis (body plan), notochord
vertebrate body plan
formation, embryo affected by yolk in egg. In mammalian, yolk to
small but have extra-embryonic structure of placenta for
nutrition
nutrition.
5) Phylotypic stage, at which they all more or less resemble each
other an show the specific features of notochord, somites and
neural tube. Fig. 2.2

5 6

The phylotypic stage Xenopus laevis: egg

At the end of gastrulation all embryos appear to be similar (the (Amphibians)
phylotypic stage).
Structures that are common to the phylotypic stage of the Advantage: easy observation, fertilized, catch (sperm, egg), low infection
vertebrates are:
1) the notochord (an early mesoderm structure along A/P axis), The egg is composed of an animal and a vegetal Animal
2) the somites (blocks of mesoderm on either side of notochord region,
i b
both
th covereddbby vitelline
it lli membrane
b ((gell
which form the muscles of the trunk & limbs), coat). Fig.2.4
3) the neural tube - ectoderm above notochord forms a tube (brain Meiosis is stopped at 1st division with apparent 1 polar
and spinal cord). body (the 2nd polar body comes after fertilization).
Box 2A
Extra- After fertilization, the cortex (the layer below plasma
embryo membrane) rotates to determine future dorsal region
nic at a position opposite to the site of sperm entry.
entry
tissue vegetal
Vertebrate embryo
to through a
phylotypic state,
but differences in
form before
Fig. 2.3
gastrulation 7 8

2
Box 2A

Cleavage of a frog egg.

9 10

Early developmental stages of Xenopus laevis Xenopus laevis : fertilization and early growth

1. one sperm enters animal region (grow to embryo, plant pore to yolk)
morula Blastula 2. completes meiosis
3. egg and sperm nuclei fuse
4 vitelline
4. it lli membrane
b lift
lifts
5. yolk rotates down (15 minutes)
6. cortical rotation occurs (60 minutes).
囊胚
7. 1st cleavage occurs (90 mins) Animal / Vegetal (A/V)
8. Every 20 mins, one cleavage
2.5 hpf
p 3.5 hpf
p 5 hpf
p 10 hpf
p 9. 2nd cleavage (110 mins) A/V 90 degrees to 1st
10. 3rd cleavage (130 mins) equatorial (4 small animal and 4 large
blastocoel -
vegetal= 8 , it is blastomeres).
11. Continued cleavage → blastomeres ↓, cells at vegetal region
hpf: hours post-fertilization large than those at the animal region.

11 12

3
 Xenopus laevis: blastulation

The blastula (after 12 divisions)

has radial symmetry.
The marginal zone will become
mesoderm and endoderm.
Marginal zone, the belt of tissue
around the equator , plays a
crucial part in future
development.
Internalization of the mesoderm
and endoderm starts at the
blastopore.

Fig 2.3 Life cycle of the frog Xenopus laevis.

In blastula stage, it is in the form of a hollow sphere with radial symmetry
13 14

Types of cell movement during gastrulation Xenopus laevis: gastrulation

Gastrulation step:
1. Mesoderm and endoderm converge and begin to move inwards at dorsal lip of the
blastopore.
2. Mesoderm and endoderm extend in along A/P axis.
3. Ectoderm spreads to cover embryo (epiboly).
4. Dorsal endoderm separates mesoderm from the space between the yolk cells, the
archenteron (future gut). Do not forget, mesoderm come from ectoderm
5
5. Lateral mesoderm spread to cover inside of archenteron
archenteron.
6. dorsal mesoderm is beneath dorsal ectoderm
7. mesoderm spread to cover gut
8. epiboly - ectoderm covers embryo
9. yolk cells are internalized (food source),
dorsal mesoderm develops into
a) notochord (rod along dorsal midline) and
b) somites (segmented blocks of mesoderm along notochord).
Invagination
Blastopore
Involution ↓
Ingression Archenteron
↓
Delamination Large
Eiboly: ectoderm covers embryo ↓
Blastocoel
↓
Close
↓
15 gut 16

4
 Xenopus laevis: Neurulation

• Neuralation or neural tube formation:

1) The neural plate is the ectoderm located above notochord and
somites.
2) The edge of the neural plate forms neural folds which rise
towards midline.
3) The folds fuse to form neural tube.
4) The neural tube sinks below epidermis.
• The anterior neural tube becomes brain. Mid and posterior
neural tube becomes spinal cord.

Gastrulation → neurulation → neural plate → fold → tube

notochord
Neural crest cell
Anterior posterior
Autonomic nerves ↓ ↓
17 Brain spinal cord
18

Fig. 2.7 Neurulation in amphibian

Xenopus laevis: Somites
The somites formation, after neurulation
The dorsal part of somites have ready begun to differentiate into dermatome
(future dermis).
The rest of each somite becomes vertebrae and trunk muscles (and limbs).
Lateral plate mesoderm becomes heart, kidney, gonads and gut muscles.
V
Ventrall mesoderm
d b
becomes bl
blood-forming
df i tissues.
i
Also at this stage, the endoderm gives rise to the lining of the gut, liver &
lungs.
Brain and spinal

Fig. 2.8 A cross-section

through a stage 22 Xenopus
embryo just after gastrulation
and neurlation are completed

Notochord begins to form in the midline

Neural plate develops neural folds 19 20

5
 The major lineages of the mesoderm Xenopus laevis: tail bud stage

• After gastrulation comes the early tail bud stage

In the anterior embryo:
a) the brain is divided,
b) eyes and ears form,
c) 3 branchial arches form (anterior arch later becomes the jaw.

In the posterior embryo, the tail is formed last from dorsal lip of
blastopore by extension of notochord, somites and neural tube.

Circulatory body cavity Fig. 2

Fi 2.9
9 Th
The early
l ttailbud
ilb d
system stage of Xenopus embryo
Scler
Myo
tome Cartilage skeletal dermis

21 22

Schematic representation of neural crest formation

Xenopus laevis : neural crest cells (in chick embryo)

Neural folds meet and adhere

Neural crest cells come from the edges of the neural folds after neural
tube fusion. Neural crest cells can form from the dorsal side of the Cells at this junction form neural crest
closed neural tube
Neural crest cells detach and migrate as single cells between the
mesodermal tissues to become:
1) sensory and autonomic nervous systems
2) skull
3) pigment cells
Closure not simultaneous
4) Cartilage → bone

Only vertebrate
Cell adhesion molecular expressed dependent
Closed tube detaches – change
Epidermal and neural plate/tube interactions may generate crest cells
in adhesion molecule
23 expression 24

6
 Zebrafish (Danio rerio
rerio)) -- A Vertebrate Model

•It is 3 cm long

•Short generation time

•Large clutch size

•External fertilization

•Transparent embryos

•Rapid development

http://zfin.org/ and
http://www.nih.gov/science/models/zebrafish/
25 26

Sphere

29h

48h
27 28

7
 Fish (Zebrafish) embryo:
•Human disease model
•Transgenics
•Reverse genetics tool

Fig. 2.26

29 30

The development of Zebrafish Characterization of Fish embryo

Telolecithal: most of the egg cell is occupied by yolk

Zebrafish Meroblastic: the cell divisions not completely divide the egg
development occurs Discoidal: since only the blastodisc becomes the embryo, this type of
very rapidly
rapidly. In 24 hr meroblastic cleavage is call discoidal
discoidal.
hours of
embryogenesis, Cleavage can take place only in the blastodisc, a thin region of yolk free
shown here, the 1 cytoplasm at the animal pole of the egg.
cell zygote becomes
into a vertebrate
embryo with a
tadpole-like form.

Fig. 2.27 Cleavage of the zebrafish embryo

31 32

8
 Fish embryo: blastula stage
Three cell populations:
At about the 10th cell division -- the onset of the About 10 cell division, the onset of mid-blastula transition: gene transcription
MBT begins, divisions slow and cell move. And formed three distinct cell
mid-blastula transition populations:
1. Yolk syncytial layer (YSL) (1)YSL (yolk syncytial layer): location of vegetal edge of the blastoderm and
2. Deep cells -- forming the embryos proper fusion produces a ring of nuclei within the part of the yolk cell cytoplasm
3 Envelope
3. E l layers
l (EVL) -- forming
f i the
th epidermal
id l that just beneath the blastoderm
blastoderm. It is important for directing some of the
cell movement of gastrulation.
ANIMAL BODY Internal YSL: the yolk syncytial nuclei move under the blastoderm
External YSL: some cell move vegetally, stay ahead of the blastoderm
margin
(2)Enveloping layer (EVL):
Made up of the most superficial
cell from the blastoderm, which
form an epithelial sheet a
single cell layer thick.

(3) Deep cells

Blastoderm Both YSL and EVL are the deep
cells, that give rise to the embryo
4 hpf: hours post-fertilization 33
proper. 34

Fish embryo: gastrulation

The fate map of the deep cells after mixing has stopped

The blastoderm
at 30%
completion of
Internal epiboly (4.8 hr)
YSL

This stage, no mesoderm, ectoderm

The fate of the early blastoderm cells are not determined. After much cell
mixing during cleavage

35 36

9
 Types of cell movement during gastrulation

Close-up of the
marginal region
Formation of the
hypoblast, either by
involution of cells at the Invagination
margin
g of the epibolizing
p g Involution
balstoderm or by Ingression
delamination and Delamination
ingression of cells from Eiboly: ectoderm covers embryo
the epiblast (6hr)
The formation of germ
layers is started.
37 38

About 90% epiboly (9 hr), mesoderm Types of cell movement during gastrulation
can be seen surrounding the yolk,
between the endoderm and ectoderm

Complete gastrulation (10.3hr)

Invagination
I
Involution
l ti
Ingression
Delamination
Eiboly: ectoderm covers
embryo

39 40

10
 Fish embryo: gastrulation

Fig 2.28 Epiboly and gastrulation in the zebrafish

Mesodermal cell
After fertilization → cell cleavage → spreading out of the layer of cell Convergence and extension in the gastrula.
(
(expressed d snailil gene))
(epiboly) → upper half of the yolk become covered by a cup-shaped flank the notochord
blastoderm→ gastrulation by involution of cell → fromed a ring around
(A) Dorsal view of convergence and externsion movements during gastrulation. Epiboly
the edge of the blastoderm → involuting cell converge on the dorsal spreads the blastoderm over the yolk; involution or ingression generates the
midline to form the body of the embryo hypoblast; convergence and extension bring the hypoblast and epiblast cells to the
dorsal side to form the embryonic shield.
(B) Convergent extension of the embryo; it is show by cells expression the gene no tail
41 42
(a gene is expressed by notochord cells)

Types of cell movement during gastrulation

Invagination
Involution
Ingression
Delamination
Eiboly: ectoderm covers embryo

43 44

11
 Chick (bird) embryo: the blastodisc (blastoderm)
Chicken
The blastodisc arises through cleavage (20 hrs.).
The blastodisc can be divided into two areas:
1) the area pellucida (a light area) surrounded by
2) the area opaca (a dark ring).

犁溝

yolk

45 Fig. 2.10 46

The life cycle of the chicken (Fig.2.11)

47 48

12
 Chick (bird) embryo: the blastodisc (blastoderm) Discoidal meroblastic cleavage in a chick egg

The posterior marginal zone forms

at the junction of the area pellucida
and the area opaca and defines
the dorsal side and posterior end of
the embryo.

The hypoblast (the source of extra-

Germinal
embryonic tissues) develops as a
layer on top of yolk and develops opaca pellucida opaca
from cells from the posterior
marginal layer and the overlying
cells of the blastoderm. It come
from two sources: the posterior
marginal zone, which lies at the ectoderm
junction between the opaca and
pellucida at the posterior of the endoderm
embryo. It develop to extra-
embryonic structure and related
with epiblast.
49 50
Fig. 2.12

Primitive streak Types of cell movement during gastrulation

Formation of two-layered
blastoderm of the chick
embryo Germinal
(A,B) Primary hypoblast cells
delaminate individually to form
islands of cell beneath the
epiblast
(C) Secondary hypoblast cells
from posterior margin →
migrate beneath the epiblast
and incorporated the poly- Invagination
invagination islands → move Involution
anterior; Ingression
As the hypoblast moves Delamination
anteriorly → epiblast cell Eiboly: ectoderm covers embryo
collect at the region anterior to
Koller’s sickle to form the
primitive streak
51 52

13
 Chick embryo: the primitive streak Chick embryo: the primitive streak

The primitive streak is a slit or line on the disc which lays down the
A/P axis. (posterior) When the primitive streak reaches its greatest length (forward),
the anterior end begins to regress back to the posterior end.
Onset of gastrulation
This structure begins to form from the posterior marginal zone and Primitive streak form at posterior → forward formation → enough
extends to a point in the central region of the disc
disc. length close and regress → Hensen’s
Hensen s node → backward
Cells move towards the streak, and mesoderm and endoderm regression → formation of head, somites and notochord… (Fig.
internalize at this site. 2.14)
The anterior end of the regressing streak is known as Hensen's
Unlike amplibians, cell Node.
not only proliferation
but also growth in
size during
size,
gastrulation in bird
and mammals.

Primitive streak 53 54

Cell movement of the primitive streak of the The major lineages of the mesoderm
chick embryo

Head, somite

Circulatory body cavity

system

Scler
Myo
tome Cartilage skeletal dermis

55 56

14
 Chick embryo: gastrulation

As Hensen's Node moves toward the posterior, several structures

form behind it:
1) The head fold (from ectoderm and endoderm)
2) The notochord and somites (from mesoderm)
3) The neural tube forms above the notochord (from ectoderm)
(The anterior structures are formed first while the posterior
structures are completed last.)
4) Neural folds fuse at the dorsal midline and neural crest cells
migrate away
5) The head fold separate, gut forms and heart pieces fuse to form
heart.

57 58

Chick embryo: neurulation Fig.2.18 Development of the chick embryo

notochord

Neural plate → neural

fold → meet midline

Intermediate somites
mesoderm→ kidney
Splanchnic mesoderm
→ heat Somite star formation

13 somites

20 somites 40 somites
Hensen’s node

59 60

15
 Mouse embryo
Chick embryo: extra-embryonic structure

Amnion
and amniotic cavity
provide mechanical
protection
Chorion maintain
shell
Allantois bridge for
oxygen and waste
Vitelline vein take
nutrient form yolk to
embryo
Umbilical vein take
oxygen to embryo Fig.2.20

Egg is small, 100mm very small

Egg surrounded by protective external coat, zona
pellucida
61 62

Development of a human embryo form fertilization to implantation

Mouse embryo: fertilization

Fertilization occurs in oviduct. (Fig. 11.26)

Cleavage occurs in oviduct: 1st at 24 hours and every 12 hours after that
to form the morula (a ball of cells). (Fig. 2.21)
• Blastomere compaction happens at 8 cell stage.
• Smooth inner membranes and outer membranes are covered with
microvilli.

(c) Morula. After further cleavage

(b) Four-cell stage. Remnants of the
mitotic spindle can be seen
between the two cells that have
just completed the second
cleavage division.
divisions, the embryo is a
multicellular ball that is still
surrounded by the fertilization
envelope. The blastocoel cavity
has begun to form.
63 64

16
 Mouse embryo: In 16 cell morula →

• Cleavage partitions the cytoplasm of one large cell At ~16 cell morula, has two group cells. A small group of internal cell
– Into many smaller cells called blastomeres mass (ICM) surrounded by a large group of external (trophectoderm)
cells.
Trophectoderm: becomes extra-embryonic tissues (such as placenta).
Inner cell mass (ICM): becomes the embryo plus some extra-
embryonic tissues.
The morula (~32 cell stage) has 2 cell fates:
1) inner 8 cells (Inner Cell Mass)
2) outer ~20 cells (trophectoderm).

blastocyst

(a) Fertilized egg. Shown here is the (b) Four-cell stage. Remnants of the (c) Morula. After further cleavage (d) Blastula. A single layer of cells
zygote shortly before the first mitotic spindle can be seen divisions, the embryo is a surrounds a large blastocoel
cleavage division, surrounded between the two cells that have multicellular ball that is still cavity. Although not visible here,
by the fertilization envelope. just completed the second surrounded by the fertilization the fertilization envelope is still
The nucleus is visible in the cleavage division. envelope. The blastocoel cavity present; the embryo will soon
center. has begun to form. hatch from it and begin swimming.

65 66

Mouse embryo: blastocyst Mouse embryo: post-implantation

In the blastocyst (~3½ days), the trophectoderm and ICM are established.
Fluid is pumped in to expand cavity and increase the size of the blastocyst.
Uterine wall
blastocyst: preimplantation (3½ - 4½ days)
The surface of ICM will become the primitive endoderm while the
remaining becomes primitive ectoderm (=
( epiblast).
epiblast)
Implantation occurs. The zona pellucida is discarded and blastocyst hypoblast
attaches to uterine wall.
Development of a human embryo form fertilization to implantation

Implantation → trophoblast giant cell invade → trophoectoderm grows →

ectoplacental cone & extra-embryonic ectoderm → primitive endoderm cover
inner surface of trophectoderm → to visceral endoderm
• In the first two days post-implantation, the mural trophectoderm (cells that are not in
contact with the ECM) gives rise to polyploid trophoblast giant cells.
• The rest of trophectoderm becomes the ectoplacental cone and the extra-embryonic
ectoderm which give rise to the placenta.
• Primitive mesoderm migrates:
1) to cover inner surface of mural trophectoderm to become the parietal (腔壁) endoderm and
2) to cover egg cylinder and epiblast to become the viseral endoderm
67 • Six days after fertilization, the epiblast is cup-shaped. 68

17
 Mouse embryo: gastrulation

6½ days after fertilization:

The primitive streak forms at the start of gastrulation at the future posterior end.
(Inside cup is future dorsal side)
Cells move through the streak and spread forward and laterally between the
ectoderm and the visceral endoderm to form the mesoderm.
Later the definitive endoderm (from epiblast) will replace the visceral
Later,
endoderm.
The primitive steak first elongates, then at the anterior tip of the primitive streak,
the node forms. (The node formed from anterior → posterior)
Then notochord and somites form anterior to the node (A/P axis).
Cells migrate through mesoderm to form endoderm (gut).

Epiblast move through

the primitive streak to
give rise to the
mesoderm and
definitive endoderm.

69 70
Amnion Chorion Allantois Fig. 2.23

Mouse embryo: late embryogenesis (neurulation) Mouse embryo: final stages of gastrulation

• By 8½ days after fertilization,

1) the neural folds form at anterior and dorsal, and 1. Complex folding
2) the embryonic endoderm internalizes to form the gut. 2. Initially on the ventral surface of embryo
• 9 days after fertilization embryogenesis is complete. 3. Internalize to form the gut
4
4. Heat and liver move into their positions
5. Head becomes distinct
Fig. 2.24
6. Embryo surrounded by extra-embryonic membrane

A P

Amnion
D Chorion
Primitive streak extend→ produce Fig. 2.25
Organogenesis in the anterior part
extra-embryonic structure Neural folds formation
Allantois
→chorion, amino, allantois
The primitive streak similar to chick
(node = Hensen’s node) 71 72

18
 Diagram showing the timing of human monozygotic twinning with relation to
Formation of the notochord in the mouse
extra-embryonic membrane

Amnion Chorion Allantois

73 74

Model organism: invertebrate

Drosophila melanogaster: early embryogenesis

The Drosophila egg is the shape of a sausage .

Meroblastic (superficial) cleavage and centrolecithal
It has a micropyle at the anterior end (site of sperm entry).
With fertilization, the fusion of nuclei is followed by rapid mitotic divisions
(9 minutes) and no cytoplasmic cleavage.
A syncytium is formed (many nuclei/common cytoplasm).
After nine divisions, nuclei move to the periphery to form the syncytial
blastoderm .

Fig. 2.30 After fertilization, no cell was form, but rapid nuclear
Fig. 2.29 Life cycle of Drosophila 75 division in a cytoplasm 76

19
Box 2A Drosophila: embryogenesis

By 13 mitoses the membranes sprout to surround the nuclei to form

cells (cellular blastoderm).
~15 cells at posterior (= pole cells) are sequestered and become
the germline.
g
During first ~3 hrs large molecules such as proteins can move
between nuclei until the cellularization occurs.
Single layer of cells give rise to all tissues (syncytium ).
Gastrulation starts at ~3 hrs.
Mesoderm forms from ventral tissue, midgut from endoderm at the
anterior and posterior ends
ends, ectoderm remains on outside
outside.
During gastrulation, the ventral blastoderm (germ band), comprises
extension.
The mesodermal tube forms from ventral tissue then cells separate
and move to internal locations under the ectoderm.
77 78

Drosophila melanogaster: gastrulation

The mesoderm becomes muscle and connective tissues.

In insects, nerve cord lies ventrally (vertebrates: dorsal).
Neuroblasts form a layer between mesoderm and outer ectoderm.
midgut (anterior & posterior) grow from threads and fuse.
= anterior and posterior midgut
ectoderm becomes epidermis.
No cell division occurs during gastrulation.
Afterward, division restarts.
Future mesoderm invaginate ventral region → intrnalized tube → cell
leave tube and migrate under the ectoderm
The surface of ventral blastoderm → cell leave and form a layer
between ventral ectoderm and mesoderm

Anterior and posterior invaginate and fuse → gut

Midgut →region endoderm
Foregut and hindgut → ectodermal origin
79 80

20
 Future mesoderm Ventral view
Fig. 2.31 Gastrulation Dorsal view
invaginate ventral region →
internalized tube → cell
leave tube and migrate
under the ectoderm germline

The surface of ventral

blastoderm → cell leave
and form a layer between
ventral ectoderm and
mesoderm →nervous
system

Anterior and posterior

invaginate and fuse → gut
Midgut →region endoderm
Foregut and hindgut →
ectodermal origin 81 82

Drosophila melanogaster: segmentation

The germ band (ventral blastoderm) is main trunk region. Drosophila melanogaster: larvae
Germ band extension pushes posterior end over dorsal side.
The first signs of segmentation grooves appear to outline The larvae hatch at 24 hrs post-fertilization.
parasegments (early embryo) which give rise to segments (late
embryo). Larval structures of note include:
Segments are formed from the posterior of one parasegment and the The anterior end is the acron.
anterior of the next. (formed form posterior to anterior) The posterior end is the telson.
Along with the head, the larvae has 3 thoractic segments and 8
abdominal segments.
The ventral side of the larvae has denticle belts, alternating patches
of denticle hairs and cuticle on each segment,
segment used for
Fig. 2.32 locomotion.

There are 14 parasegments: Fig. 2.33

3 mouth, 3 thorax, 8 abdominal.
83 84

21
 Drosophila melanogaster: metamorphosis
imaginal discs
Three instar stages of larval life are separated by molts.
• 1st instar 2nd instar 3rd instar
molt molt
3rd instar larvae forms pupae (pupa) to undergo metamorphosis.
The adult tissues arise from imaginal discs and histoblasts.
imaginal discs: small sheets of epidermis (~40 cells each of cellular
blastoderm) which grow throughout larval life.
Imaginal discs: 6 leg, 2 wing, 2 haltere, 2 eye-antenna, plus genital,
head discs histoblasts
and ~10 histoblasts: nest of cells in the abdomen which give rise to the
abdominal segments.

Larval epidermis degeneration begins prior to imaginal disc eversion

Imaginal disc cells and histoblasts will replace the larval epidermis
Antenna
Formation of adult abdominal segments - gene expression in haltere
histoblasts Fig. 2.34 Imaginal discs vs. adult structure Genitalia
Imaginal discs 85 86

Caenorhabditis elegans: the model of nematode

THE WORM

After
gastrulation

In case of self-fertilization
there are ~ 0.1 - 0.3% male
worms in the population.
Fig. 2.35 Life cycle of nematode http://www.wormatlas.org/handbook/contents.htm
87 88

22
 the model of nematode Press Release: The 2002 Nobel Prize in Physiology or Medicine
7 October 2002
The Nobel Assembly at Karolinska Institutet has today decided to award
Small nematodes that are 1 mm long and 70 µm in diameter. The Nobel Prize in Physiology or Medicine for 2002 jointly to
19,000 gene
Sydney Brenner, H. Robert Horvitz and John E. Sulston
Small number of cell (558, first larval stage)
T
Transparency off embryo,
b and
d growth
th rapid
id for their discoveries concerning
The adult hermaphrodite (maless can develop) undergo rapid "genetic regulation of organ development and programmed cell
development. death"
The egg has a 50 µm diameter which forms a polar body after
fertilization, nuclear fusion occurs followed by a set pattern of
cleavage.
The normal pattern of cell division has been mapped.
Many cells undergo programmed cell death.

Hermaphrodite: 959 cells from 1090 somatic nuclei of which 131

undergo programmed cell death; 300 germ cells undergo
apoptosis; 116 of the 131 dying cells are cells of the nervous 1942
1927 1947
system and ectoderm 89 90

Molecular Regulation of Apoptosis

C. elegans

mutagenize
Non- apoptotic
apoptotic

wildtype
CED mutants
(Cell Death
abnormality)

91 92

23
 Fig. 2.36
Cleavage of
the nematode
embryo

Fertilization →polar bodies formation → asymmetric cleavage → anterior AB

cell, smaller posterior P1 cell

DIC image

Fig. 2.38 elegans larva at the L1 stage.

Fig.2.37 Anus
Cell lineage and cell fate Pharynx
in the early nematode Primordium
embryo

93 94

Invertebrate: Sea Urchin Sea Urchin: blastula formation

Radial holoblastic cleavage (isolecithal) The blastula stage of sea urchin development begins at the 128 cells.
The 4th cleavage, very different from the first three. In animal pole, four cell Blastulation: The cells form a hollow sphere surrounding a central cavity
divide to 8 blastomeres and with the same volume (the 8 cells also called (blastocoel). Every cell contact with proteinaceous fluid of the bastoceol
mesomeres). In vegetal pole, undergoes an unequal cleavage to four large (inside) and with the hyaline layer on the outside.
cells (macromeres) and four small cells (micromeres). About 9th or 10th cleavage, cells become specified and they end develop cilia.
Ciliated blastula → rotate within fertilization envelop (E→F) → vegetal pole of
The animal mesomeres divide Bastula become thicken
equatorially to produced (forming vegetal plate) →
two tiers: an1 and an2. then animal pole synthesis
The vegetal macromeres and secret hatching
divide a small cluster enzyme → digest
beneath the large tier. (not fertilization envelope →
equal) embryo is a free swimming
4th hatched blastula.
cleavage
128 cells blastula.

Meridionally

rotate
95 96

24
Fate maps and the determination of sea urchin blastomeres

Fate map of the zygote

Late blastula with ciliary tuft
and flattened vegetal plate

blastula

Fate map and cell lineage of the sea urchin.

97 98

Formation of syncytial cables by primary mesenchyme cells of sea urchin

SEM of spicules
formed by the
fusing of primary
mesenchyme
cells into syncytial
y y
cables

Gastrulation star
C: SEM of primary
mesenchyme cells enmeshed
in the extracellular matrix of
earlyy gastrula.
g
D: Gastrula-stage
Pluteus larva mesenchyme cell migration
Prism-stage larva
The extracellular matrix fibrils
of the bastocoel lie parallel to
the animal-vegetal axix
99 100

25
 Ingression of primary mesenchyme cells Invagination of the vegetal plate

CSPG release → into inner lamina → osmotic

gradient ↑→ absorb water → swell inner lamina
SEM of external surface ,but outer lamina attached does not swell →
off the
th early
l gastrula
t l inward

Fertilization envelope

101 CSPG: chondroitin sulfate proteoglycan 102

Entire sequence of Identification of developmentally important genes

gastrulation in sea urchin
The developmental genetics of Drosophila and mice are best
known.
Homologous genes identified in these organisms are found in other
species.
Dominant (or semi-dominant) mutations: one copy of mutant gene
produces mutant state. These are more easily recoginzed, they
don’t cause the eayly death of the embryo in the heterozygous.
Recessive mutations: two copies of a mutant gene gives the
mutant state.

Allele: The gene is contributed by the male and female

Homozygous: both alleles of a pair carry the mutation
Heterozygous: just one copy of the mutant gene is present

103 104

26
 Recessive mutation vs. Semi-dominant mutation

-/-

Most mutations are recessive, but

usually die in embryo.
105 106

Developmental gene can be identified by induced mutation and

screening

Genetic screening to
produced
homozygous
yg mutant
zerbrafish embryo

Heterozygous

Embryos
homozygou
s the
induced
mutation will
heterozygous
be found in
the offspring
of 25% of
the matings
107 108

27
 Mutagenesis and genetic screening strategy for identifying
developmental mutants in Dorsophila main patterns of cleavage

phylotypic stage
DTS: dominant temperature-
sensitive mutation, up 29oC Time vs. developmental events
→ death
b: a non
non-developmental
developmental lethal T
Types off cell
ll movementt during
d i gastrulation
t l ti
recessive
Primitive streak

gastrulation

Neurulation
ethyl methane sulfonate
human monozygotic twinning

Syncytium

imaginal discs and histoblasts

109 Dominant (or semi-dominant) mutations 110

28