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4-Specification I 11

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4-Specification I 11

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What Are Cell Differentiation

and Cell Specification?

• Development of differentiated cell types


– different in function, cell shape, internal
structures, proteins and biochemical pathways
• Preceded by commitment to fate
– Specification: can differentiate in neutral
environment, but reversible in new
environment
– Determination: irreversible even in new
environment
What Are the Different Modes of
Specification?
• Autonomous Specification

– Independent of other cells in embryo (self-


differentiating parts)
– Determined by internal cytoplasmic factors
– If cleavage patterns are invariant, then cell fates will be
invariant
– “Mosaic” development
– Characteristic of invertebrates
What Are the Different Modes of
Specification?

• Conditional Specification (“Non-Autonomous”)

– Depends on conditions (cell interactions)


– More variable (variations in which cells are assigned
which fates)
– Capacity to “regulate”
– Characteristic of vertebrates (and some invertebrates)
What Are the Different Modes of
Specification?
• Syncitial Specification

– Interactions between regions of cells before


cellularization
– Cells have no specific cell fate before cellularization
– After cellularization, mostly conditional specification
– Most insects
Chabry’s Experiment 1887

• Mollusc
• Certain cells
form trochoblasts
(ciliated)
• In isolation
• autonomous
Roux’s Frog Cell Destruction
Experiments (1888)
• Defect experiment
• Frog embryos
• One blastomere makes
only half of embryo
• Support for
autonomous
specification?
Tunicate Specification is Mostly
Autonomous
• Tunicate embryo

– Isolation experiment
– Separate 8 cell stage
quadrants
– Each forms only what
it would have formed
and does so
autonomously
Cytoplasm Contains Determinants
Cytoplasmic Autonomy Extends
to the Molecular Level
• Tunicate Embryo
(1977)

– Isolated muscle forming


cell lineage makes
muscle
acetylcholinesterase
– If cytoplasm injected
into isolated b4.2
causes it to make the
enzyme
Driesch’s Sea Urchin Isolation
Experiments: Regulation (1892)

• Sea urchin
• Each blastomere
gives rise to
complete larva
• Supports
conditional
specification
Conditional Specification in Frog
Transplantation Experiment
• Cell fate according to
new environment (A)
– fate not fixed
• Embryo makes up for
what was taken away
(B)
– regulation
Gradient of Fate-Determining
Molecules in a Syncytial Embryo
Stem Cells and Commitment
• Cells that divide to form one copy of itself and one
different
– Pluripotent stem cells can lead to many cell types
– Committed stem cells can lead to fewer types
– Progenitor cells committed to one or few cell fates: not
stem cells
• Early cells have most potential (embryonic, fetal)
Restriction of Potency with Time
• Committed cells usually don’t change in
new environment
• Blood cell lineage:
Mutants Identify Some Fate-
determining Molecules
• What would be effect of a mutation in a
fate-determining molecule?

• These molecules must be acting very early


– Packaged by the mother
– Mutation in gene will influence the mother’s ability to
make viable embryos
– “Maternal-effect” genes

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