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Animal Developmental Biology - Lecture notes - Lecture 1

Animal Developmental Biology (University of Leeds)

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Animal Developmental Biology

Lecture 1

1. Fertilization  Zygote (single cell) Animal Development  Adult

2. Model systems (easy to study, yet representative of animal development

more generally)
a. The frog
b. The fish
c. The chick
d. The mouse
e. The worm
f. The fly

 Conjoint twins
 Distinct from the shoulders up
 Died 8 months old

 Only adults in America joined at the head, 42 years old

 Pennsylvania
 Share bone, tissue and blood lines, joined at corner of left eye
 Share brain tissue, but have individual identities
 Siamese twins
 Comes from two siam twins that had joined livers
 Proclaimed their oneness
 Seen as having political or religious significance
 Signify political union…
 Conjoined twins as a natural phenomenon

 The heart apex is pointing down to the right

 Spleen is on the right
 Stomach is on the right
 All of ours are on the left
 Cytosinversis (left to right reversal of the internal organs; rare but in
conjoined twins)
 Seen in Rita and Christina

 13 days after conception, embryo begins to organise itself (gastrulation)

 Gives the embryo its geometry
 Remove the membrane surrounding the egg, to it upside down, and excise
the tissue, then graft into another embryo on the other side
 Would the transplanted cells behave differently from the hosts?
 Almost all of the embryos died
 Transplanted cells organised the host cells into an extra, but conjoined
creature
 Individual cells don’t govern their own fates
 Organiser limited the potential of cells around it, was the origin of order

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 Organiser = clump of cells

 Amsterdam = museum with collection of mutants

 Terrontolity? – study of monsters
 Most of the specimens in the museum are mutants,

 Mermaid syndrome
 Legs are fused together, feet protrude like flippers
 Usually fatal because it affects many organs, like the heart
 Midline disorder, lower half of body collapses in on itself
 Operations took plastic surgery to a new level, lower half of body had to
be completely rebuilt
 Tiffany, only person in the world to survive with it to the age of 16

 Causes of it have been obscure, but knew it was the organiser

 Needed to find the substance secreted by cells in the organiser
 Discovery was called Corden – it was a signalling molecule
 Retinoic acid

 If you cut the tale off a tadpole it regenerates, but if you put it in retinoic
acid you get a tangle of legs

 Last body part to form – Face

 Pig that has 2 faces, 2 snouts, 4 nostrils, 3 eyes (Called Ditto)
 Had an optic nerve that connected to the third eye, but unsure if he could
see through it
 Chicken with 2 upper beaks
 Sonic gene – place a soaked bead in a different area in the embryo, but
still in the face region


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Lecture 2

 Xenopus
o Page 43 in Gilbert
o Red part is the Deuterostomes which include the vertebrates

 Deuterostomes

 Page 27
 Fertilization (0h. 1 cell)  Cleavage (rapid cell divison = 6h. 10,000 cells) 
Gastrulation (re-arrangement of cells from interior to exterior, generates 3 germ
layers; 10h. 30,000 cells)  Organogenesis/Neurulation (generates the organs,
infold of cells from the dorsal surface to give rise to the CNS; 19h. 80,000 cells)
 Larval stages (110h. 1,000,000 cells)  Maturity  …

 Cell division occurs precisely from the animal pole at the top, bottom is the
vegetal pole?
o Can be seen clearly as its pigmented
o Higher concentration of yolk at vegetal pole
o Yolk inhibits cytokinesis (division of the cell)
 Cleavage furrow moves down through the embryo to reach the vegetal pole
 Second cell division starts to take place, when first division is still incomplete
 Third cell division starts before second has finished, at right angles to first 2
divisions; embryo now divided up into 8 cells
o 4 at the animal pole, 4 at the vegetal pole
o Smaller cells at the animal pole and larger cells at the vegetal pole
 Blastula is the final stage
o 10s of thousands of cells, with fluid filled cavity (blastocoel) towards
animal pole
 Individual cells referred to through these stages are blastomeres
 Cells of the animal pole will give rise to the ectoderm, cells around vegetal pole
gives rise to endoderm, and cells around equatorial region give rise to the
mesoderm

 Cell division occur every 30 mins, very rapid

 Single cell zygote is over 1mm across, 1000 larger then a bacteria
 Genomes that need to be duplicated are so much larger than in the bacteria
 The embryo never grows in diameter, it just undergoes organisation
 Each cell generated all has its own copy of the genes

 Blastula then undergoes gastrulation

 Mid-blastular transition – when cell division stops and cells start to re-arrange =
gastrulation
 The cells start to fold into the interior from the dorsal surface (back of the
animal)
o Cells left on outside are the dorsal-blastopore lip?
 Ectoderm gives rise to the outside surface, endoderm will line the inside/gut of
the animal, mesoderm fills the middle
 Archenteron = fluid filled cavity  Gives rise to the lumen of the gut
 As it folds in, the mesoderm forms between the 2
 Ectoderm cells move around the outside at the endoderm folds in
 Few endoderm cells still left on the outside, which is the yolk plug
o ….

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 As the endoderm folds in, it reaches the other side of the embryo, and where it
comes into contact with the ectoderm with no mesoderm = where the mouth
forms (secondarily)
 Anus forms at the connection between the archenteron and …
 Deuterostome comes from
o Means mouth forming secondarily

 Neurulation
o To generate the CNS
 Small circle is the notochord, mesodermal tissue, runs from head to tail just
below the surface
o Send signals to overlying ectoderm that they’re going to give rise to the
CNS
o That region of the ectoderm are going to fold into the interior, which
generates a neural groove as the cells fold up and close over
o To pinch off the neural tube running all the way down the back

 Neuropore closure failure

o The ends cant be sealed up
o Not only in the frog, but in us as well
o The ends are referred to as Neuropores
o Has serious consequences (exposes brain the outside service so it
degenerates = anencephaly; if tail end fails to close = spina bifila)

 Vertebrate cloning was achieved a long time ago

 King, 1952
o Blastula stage nuclei in Rana pipiens
o Transferred those to anucelated zygotes (nucleus removed)
o Embryo would develop fine through to adult stage
 Gurdon, 1962
o Differentiated tadpole intestinal nuclei in Xenopus laevis
o Low success of the experiment
o But could be increased by …
o Shows a fully differentiated cell could be used to create a new individual,
and could generate all the different cell types the animal needed  The
information was still retained even after it was fully differentiated

 Fate Maps
o A map of the fates of the cells that are present in the embryo at a
particular time
o Glastular stage (label cells on outside using a dye, follow through the
development, and can see what cells are derived from the previously
labelled cells)

 Highly regulative development

 If placed on the ventral side (belly rather than back), then the planted cell gives
off signals telling the embryo…
 …
 Second embryo that formed is largely composed of cells from the host…

 1-cell stage
o up to 80 mins

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 2-cell, 4-cell, 8-cell stages

o up to 25 mins
 As they change through the cell cycle, …

 Embryos are large, accessible (external fertilisation), robust and well-studied

o Signalling molecules that direct development have been biochemically
purified (KEY THING)
 But Xenopus laevis is an allotetraploid as a result of an evolutionarily recent
hybridization, making the species far from ideal fro genetic approaches
 The advances in the field were made through a genetic approach
 Allotetraploid = recently during its evolutionary history, 2 species have come
together and hybridized to generate this species
o Its effectively a tetraploid
 Genetics is hard enough with a diploid

 Xenopus tropicalis is the only one in Xenopus species which is diploid

o The adult is much smaller than X. laevis
o Generation time is 4-6 months
o Transgenesis procedures developed
o Genome sequence completed 2010

 Desert tadpole mutants in tropicalis

o Lack of circulating blood
o Heart is there but no blood to pump
o The blood cells are moving around in the tail of the wild type

 Key points
o Classic model, very important in developmental biology
o Robust to experimental manipulation
o Identification of inducing signals
o

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Lecture 4 REVISION

Disadvantages of mammals as model organisms for studying development

 In viviparous mammals, embryonic development is entirely in utero
 Genetic manipulation is more difficult
 Long gestation and generation times
 Ethical restrictions
Advantage! – It’s a vertebrate mammal so closely related to use

The sperm must fuse with the egg

 Receptors on the surface of the sperm, bind specifically to the zona
pellucida protein 3 (ZP3)
 Binding triggers the acrosome reaction (releases enzymes from the head
of the sperm)
 Leading to digestion of the zona pellucida and then membrane fusion

Mechanisms operate to prevent polyspermy (fusion of more than one sperm

with an egg)
 Fertilization has occurred with 2 sperm cells fusing with an egg
 18 chromosomes from each sperm nuclei
 Sperm centrioles are brought into the cell at fertilization (2 centrioles);
they divide so we have 4 centrioles
 These centrioles organise the spindle of cell division
 Each centriole sets up the corner for the spindle
 6 spindles would be set up, so 6 equators for the cells to form
 Each cell has an inappropriate number of chromosomes, so development
would fail

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
 Mammals – sperm fusion leads to the release of the contents of cortical
granules from the egg surface
 These granules contain enzymes that modify ZP1, 2 and 3; so sperm cells
no longer penetrate the zona pellucida
 In other animals the cortical granules lead to the lifting of the vitelline
membrane (tight against the surface of the egg) from the egg surface
forming the fertilization membrane
 Both of these aspects are very slow!
 When sperm fuses with egg, its initially a fast but temporary fusion
 So a fast but temporary block to sperm fusion comes from an immediate
change in the electric potential of the cell membrane

Movement of pre-implantation embryo down the maternal reproductive tract


 Oocyte is released from the ovary into the oviduct (where fertilization
occurs)
 Initial cleavage occurs during this process
 It reaches the uterus where it implants itself – subsequent development
occurs

Mammalian cleavage is equal, asynchronous, rotational and slow

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 Equal
o The initially formed cells are equivalent in terms of developmental
potential (can contribute to any of the cells subsequently formed)
 Asynchronous
o Don’t just et 1,2,4,8,16.. stagesl you also get 3,5,7,9 stages
 Rotational
o First cell division occurs in one direction, whereas in the next cell
division, the next 2 cells both divide at right angles to the first
stage, but also at right angles to each other
 Slow
o Takes 24 hours to part

Compaction occurs at the 8 cell stage


 Compaction = they appear to become stuck together
 Tight and gap junctions are forming between the cells
 Tight junctions (important for sealing up the embryo), generate an inside
and outside
 Gap junctions (important for communication), allow small molecules to
flow between cells
 Cell division carries on and fluid filled cavity forms (blastocoel)

Differentiation of the inner cell mass and the trophoblast, and development of
the blastocoel, to form the blastocyst

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
 This stage is a blastocyst, not a blastular
 Consists of essentially 2 cell types
o Outside cells (trophoblast or integrin proteins) – generate the
placenta for interaction with the mother
o Inside cells (inner cell mass or hypoblast) – give rise to the foetus

Primary cell lineages in the mammalian embryo


 Primitive streak – when the cells moves into the inside

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The blastocyst has to hatch from the zona pellucida before implantation
 Blastocyst produces enzymes that digest the ZP and allow it to hatch out
 Important event that has to be carefully controlled
 If escapes too early, will result in eptopic pregnancy (implant into any
tissue its up against)

The hatched blastocyst then implants into the wall of the uterus
 The inner cell mass now consists of the epiblast and the hypoblast
 After implantation, a second fluid filled cavity begins to form (amniotic
cavity)
 Forms on the inner side of the cell mast where the epiblast is

The amniotic cavity grows

 The extra embryonic endoderm forms the yolk sac (although there is no
yolk in the mammalian embryo)

Gastrulation
 Occurs with cells moving inwards from what will be the dorsal surface,
forming the endoderm then the mesoderm (around 16 days)
 The node (equivalent of the frog organiser) is seen at the end of the
primitive groove

Human neurulation, with closure of the neural tube at the dorsal surface

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
 The amniotic cavity wraps around the entire embryo, with the ectoderm,
mesoderm and endoderm folding round to meet on the ventral midline

Neural crest cells (characteristic of vertebrates)

 The last cells derived from ectoderm to move into the interior
 Migrate individually to various locations forming the: cartilage, bone and
muscle of the face, the peripheral nervous system, the pigment cells of the
skin and adrenal medulla (modified sympathetic ganglion – in adrenal
gland above kidney)
 Equivalent to neural crest cells have been seen in other chordates in the
past year
 The shape of the face arises from neural crest cells that have migrated
from the dorsal surface of the embryo

Formation of the amniotic cavity in the chick

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 For birds, the chorion is important for gaseous exchange

 In mammals, its important for the interaction in the placenta with the
mother
 Evolution of these events was crucial, allowed animals to escape the need
to develop in water and invade the land
 Allantoic membrane is where toxic products develop away from the
embryo

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Lecture 5 REVISION

Mammalian cells are very highly regulated

 The cells of the inner cell mass of the blastocyst show totipotency (ability
to differentiate into any type of body cell)
 Particularly regulative due to the nature of the early development
implanting in the uterus wall

1. Origins of identical human twins (1 in 400 births)


 Result of a single fertilization event generating a zygote that divides into 2
individuals
 3 different scenarios can occur
 A – if separation occurs before day 5 of fertilization (1/3 of instances);
end up with 2 blastocysts that implant into uterus wall, and develop 2
placentae
 B – between day 5 and 9 (2/3 of cases); single blastocyst but inner cell
mass separated into 2 clumps of cells, single placenta generated, shared
by 2 individuals; develop in own amniotic cavity, so effectively develop in
own environment
 C – separation occurs after day 9 (very rare); derive from a single
blastocyst, single clump; embryos develop sharing the amniotic cavity and
risk being conjoined = developing in same fluid filled environment, cells
can become intermingled, but development carries on as best it can
 Conjoined twins are joined through the same part of their anatomy
 Hagness (the wrong way) of the organs within twins, as well as the whorl
on the head
 One twin has normal hagness the other has opposite hagness (left-right
orientation)

2. Cloning by embryo splitting – here in a non-human primate (Rhesus)

 Possibly derive a clone of individuals from early on in embryos
 Take an 8-cell mammalian embryo split into 4 pairs of blastomeres, which
can be replaced in empty zonae for culture

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 Culture in vitro for a while, then implanted into a surrogate mother and
will develop due to its high regulative nature
 A miscarried pregnancy and normal development resulted from transfer
of two quadruplet embryos each to two surrogates

3. PGD – Pre-implantation Genetic Diagnosis

 Cell is removed from an early human embryo for genetic testing
 The remaining cells will give rise to a complete individual
 After genetic analysis of the isolated cell, the remaining embryo is
returned to the mother with implantation

4. Formation of chimaeras
 Cells from different mouse blastocysts can be mixed together and all can
contribute to all tissues


 Once the 2 embryos have been mixed up, the cells can still pass signals
between each other and generate a single blastocyst
 Place into a surrogate mother and will develop fine
 Chimaera – ½ cells from one strain, ½ from the other (hence 2 tone
mouse)
 Cells from multiple 4-cell Rhesus monkey embryos can be mixed together
to create chimeric infants

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
 Single chimeric individuals produced

5. ES cells – embryonic stem cells

 Inner cell mass can give rise to any cell types present in embryo
 They can be isolated then cultured, as ES cells
 These cells divide and are maintained in an undifferentiated state
 By applying the right protein signals, these cells can be directed to
differentiate into different cell types (e.g. blood cells, nerve cells, muscle
cells), potentially for medical applications

6. Reverse genetic analysis in the mouse

 ES cells can be genetically manipulated in vitro
 Once these manipulated cells are returned to the host blastocyst, they can
contribute to the germ line of the embryo and so genetically manipulated
strains can be established
 These cells can be maintained in culture in large numbers
 Manipulate cells and change their DNA content
o Indicated in the picture below


 Can use electric shock to electro berate the DNA into these cells
 This process ^^^ allows us to generate a transformed strain of mice
 The embryo will be chimeric, consisting of cells from host blastocyst and
ES cells in culture

DNA introduced into ES cells can recombine homologously with the cell’s own
genetic material

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 DNA at the top is being introduced, contains an extra bit of DNA in red
 Now it can recombine with the chromosomal copy of the gene
 When recombination occurs the extra DNA is inserted into the
chromosomal copy of the target gene
 Can select for the very rare events, and identify the specific cells where
the event has occurred
 If inserted randomly, the blue bit of DNA will be inserted as well
 Can generate a strain of mouse that has a very precise modification

2007 Nobel Prize for Physiology or Medicine

 Capecchi, Evans and Smithies
 “Principles for introducing specific gene modifications in mice by the use
of embryonic stem cells”

Reverse genetics in the mouse to knockout sets of genes


 Top  wild type; Middle  hox10aaccdd knockout (lumbar to thoracic
vertebrae); Bottom  hox11aaccdd knockout (sacral to lumbar
vertebrae)
 Hox genes originally identified in D. melanogaster
 Hox10 exists in 3 different versions (a,c,d)
 If you knockout one of these versions, it doesn’t have an affect
 Need to inactivate all 3 copies if you want to see an affect
 Effectively 6 copies though (animal is diploid)

7. Induced Pluripotent Stem (iPS) Cells

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
 Transcription factors responsible fro the pluripotent state in cells in the
embryo can induce pluripotency in adult cells
 Possible because inner cell mass cells have totipotency
 This power could be provided by just 4 transcription factors
 Could confer totipotency on cells of which these 4 TF’s were introduced
 induced pluripotent stem cels (iPS)

The Nobel Prize in Physiology or Medicine 2012

 Gurdon and Yamanaka
 “for the discovery that mature cells can be re-programmed to become
pluripotent”

CRISPR-Cas9 mediated correction of a mouse model of Duchenne Muscular

Dystrophy
 Genomic correction was made with injection of nucleic acid into the
zygote

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Amphioxus (or lancelot)

 e.g. Branchiostoma lanceolatum
 Not a mammal or vertebrae
 Chordates, but don’t have a backbone
 Notochord instead, found below the dorsal surface and provides rigidity
 Dorsal neural tube
 Muscles running the length of the body


 Early stages of development are like those for the sea urchin
 Later stages are more vertebrate like


 The cephalochordates and urochordates (invertebrates) branched off
during evolution, before the two whole genome duplication events
leading to the vertebrates
 Cephalochordates and urochordates don’t have extra copies of genes so
easier to study


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Lecture 6 REVISION

Sea squirts are urochordates (ascidians), in the chordate phylum

The tadpole-like larval stage of the ascidians demonstrates the affinity with
vertebrates
 Has a dorsal nerve chord like us


 Green cells = notochord (provides strength in tail)
 Can see individual cells and nuclei – indicates simplicity of the animal

In some species of tunicate, different areas of cytoplasm in the egg are different
colours and segregate to different tissue types during development

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 Animal pole where the oocyte nucleus is

 Vegetal pole where there is a small amount of yolk
 The different colours appear to reveal cell-intrinsic determinants
(directing them along different fates)


 After fertilization, cleavage occurs as well as cell divisions which are at
right angles to one another
 Through the animal and vegetal pole, generating a left and right side with
perfect symmetry
 4 cells at animal pole, 4 cells at vegetal pole
 Radial cleavage is characteristic of dueterostomes
 Around 32-64 cells, gastrulation occurs forming the 3 germ layers

Classic view
 Ascidians
o Mosaic or determinate development
 Vertebrates
o Regulative or indeterminate development
 The way in which the cytoplasm is divided up, looks as if something inside
it is directing it
 Mosaic development = depends on what is within them, not through
communication between cells

After separation at the 8-cell stage, the cells continue development as in the
intact embryo  determinate development

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
 An 8-cell stage embryo  Divide it up into 4 pairs of cells and separate
them  Pairs of cells from animal pole will develop into ectoderm; cells
from vegetal pole will give rise to endoderm and mesoderm (muscle and
notochord)
 Different from mammals! (Each pair would give rise to complete embryo,
not a particular germ layer!)
 Notochord sends signals to the overlying ectoderm to direct nervous
system development

Currently emerging view

 Early ascidian development actually depends extensively on intercellular
signalling, but these are identical in all embryos!

Similarity of Ascidian and Amphibian fate maps

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
 Top are Ascidians
 Can see the same relative positions of different cell types

Ciona savignyi (solitary sea squirt)– A real genetic system for the chordates
 Advantages
o Hermaphrodites with cross and self-fertilization
o Small genome – 1.8x108bp ~ 15,500 genes; already sequenced in
2005
o Genes are present in single copy per haploid genome
o Simple anatomy – 2,500 cells at hatching
o Cheap and easy to maintain – filter feeders; just need access to
large bodies of sea water
 Disadvantages
o Marine labs only
o Generation time is 8-10 weeks
o Seasonal – Jan-Mar poor growth, although gametes throughout the
year
o Size – 200 adults in 15 litres, although larvae only 1mm long
 Or Ciona intestinalis
o Genome sequenced in 2002
o Transformation of hundreds of embryos at a time by
electroporation

Notochord specification
 Brachury (or T) fused to GFP
 Brachury/T has a primary role in Notochord specification in all chordates

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 Human T (brachury) gene duplication confers major susceptibility to

familial chordoma
 This gene comes on once a cell during development has been committed
to the notochord
 This is the ancestral function of this gene

Notochord development
 Bra switches on other genes, starts a cascade of gene expression
 Genes expressed in the early phase have multiple binding sites
 Middle expressed genes have a single binding site at the end of them
 Last stage, the genes have no binding sites for Brachury
o But their expression depends on other genes

Elongation of the tail is driven by a change in the shape of the notochord cells
with no further cell division
 They now look square (but are actually cylindrical)
 Their width has shrunk but length has increased

Myosin and actin


 Myosin and actin are organised in each cell of the notochord to drive
elongation, by reducing the cell diameter with no change in cell volume,
upon contraction
 As the actin structure constricts, the cell stretches along anterior-
posterior direction

The mechanism for notochord elongation is derived from apparatus driving

cytokinesis
 Previously identified for elongation of the C. elegans embryo
 A ring constricting down to separate the cells into 2, without actual cell
division

The head of the larval stage has 2 pigmented eyespots

 The otolith senses gravity and the ocellus senses light
 Abitua et el. Suggested that the cell lineage giving rise to these pigmented
cells in this invertebrate may be the rudimentary neural crest (that move
in at the end of neuralation)
 The pigment cells in our skin arise from the neural crest, a very important
vertebrate specific tissue

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 Evolution of the neural crest is the to be key to the switch from filter
feeding to active predation in ancestral vertebrates
o Also, because they give rise to all the cells in our head


 Black pigmentation stands out clearly against transparent larval cells

Sea Urchins

Still in the deuterostomes, but no longer chordates

 Echinodermata

Cleavage patterns in sea urchins


 Initial cells division gives rise to oral (left cells), right (top cell), left
(bottom cell) and aboral (right cell) cells
 4 cells above another 4 cells
 Divide in different fashion
o Animal pole cells divide, followed by vegetal pole cells

Gastrulation and morphogenesis to the pluteus larva

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
 Single cell layer thick
 Fluid filled cavity
 Hatches out and the cilia around the outside allow it to move
 Gastrulation starts on the vegetal side
 Endoderm moves inside
 Blastopore gives rise to the gut as it folds into the interior
 Mouth forms when the endoderm and ectoderm come into contact
o Still formed secondarily (because it’s a deuterostome)

Show regulative development


 All of the different tissues making it up are formed from each of these
cells
 Signals communicated between these cells to make sure all the structures
needed in this embryo are generated

Ripe sea urchins can be induced to spawn – yielding huge quantities of gametes
 Separate sexes for sea urchins
 Mix large numbers of eggs and sperm, and they will be fertilized at the
same time and will undergo synchronous development

The 2001 Nobel Prize for Physiology or Medicine

 Hartwell, Hunt, Nurse
 “Key regulators of the cell cycle” – started with cleavage in the sea urchin

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 lOMoARcPSD|4349629

Lecture 8 REVISION

Life cycle of Drosophila

 9 days from fertilization to mature adult


 Life cycle of 9 days means we can do experiments quickly (but slower
then C. elegans, faster then other vertebrate model systems)

Stages of Drosophila embryogenesis

 Mature egg is not round, its elongated
 Anterior to the left, posterior to the right (apparent because of the
flattened top of the egg, which reflects the second axis of the animal which
will develop = dorsal surface)  curved surface = ventral surface
 After fertilization its just a single cell
 The egg is very yolk rich = greatly inhibits cytokinesis


 Superficial cleavage (1 – 2)
o Nucleus will undergo mitosis repeatedly, with no cytokinesis at all
o Nuclear divisions are 10 mins apart = stage 2 (512 nuclei in a
single cell)

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 Most nuclei move to the outside (stage 3)

 Syncytial blastoderm (3 – 4)
o A few nuclei at the posterior pole gives rise to the pole cells, and
the membrane at the outside moves in and around these 5 cells to
form a separate nuclei each
o They give rise to the germ line and the gametes
o

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