Entwickslungmechanik

Developmental Mechanisms
Cell Specification
Autonomous
Conditional
Syncytial
Cell Specification
 Differentiation
 The process and the processes associated
with a cell becoming specialized
 Occurs in multiple steps
Cell Specification
 Commitment
 Specification

 Determination

 Terminal differentiation
Cell Specification
Commitment
 Specification
 A cell is said to be specified when:
 Cells differentiate autonomously when removed
from normal environment (embryo) and placed in
a neutral environment (culture medium)
 Placing cells into a non-neutral environment (a
different place in the embryo) will cause the cells
to follow the fate of other cells the new location
rather than their original fate
Cell Specification
Commitment
 Determination
A cell is said to be determined when:
 Cells
differentiate autonomously even
when placed in a non-neutral environment
 When moved to a different location within
the embryo, the transplanted cells
differentiate according to their original fate
Cell Specification
Terminal Differentiation
 When a cell can no longer change or be
changed into anything other than the cell type
it is
 Can be associated with permanent changes
in DNA
 Methylation is a prominent factor
 B-cells (plasma cells) rearrange the
immunoglobulin (Ig) genes so that they can now
only form a single type of Ig
Autonomous Cell Specification
 Characteristic of most invertebrates
 Differential acquisition of cytoplasmic
determinants (morphogenetic factors)
 Invariant cleavages and invariant
lineages
 Gives rise to mosaic development
 Specification precedes cell migrations
Autonomous Cell Specification
 Isolated blastomeres will differentiate
into the same cell type they would have
differentiated into if left in the embryo
 Removal of cells from the embryo
results in an embryo missing the cell
types the removed cells would have
differentiated into
 Embryois in essence a mosaic of cells.
Removal of any cell from the early
embryo results in a missing region of
the embryo
Autonomous Cell Specification
Autonomous Specification in Mollusc Embryos
Autonomous Cell Specification
 Autonomous
differentiation of
dissociated
blastomeres of
tunicate embryo
Autonomous Cell Specification
 Autonomous specification implies
localized morphogenetic determinants
 Morphogenetic determinants become
partitioned into individual blastomeres
during cleavage stages
 Morphogenetic determinants are both
proteins and mRNAs
Conditional Cell Specification
 Characteristic of all deuterostomes
 Specification via cell-cell interactions
 Cleavages are variable – no definite cell
lineages
 Development is regulative
 Cell rearrangements precede &
accompany specification events
Conditional Cell Specification

Transplantation experiment Defect experiment

Conditional Cell Specification
 Conditional specification implies the
interaction of cells in some manner
 Cells must recognize their neighbors
 Cells must influence each others
differentiation
Conditional Cell Specification
 Cell-Cell interactions
 Restriction of fates
 Inhibition by neighboring cells
 Inductions
 Neighboring cells initiate induction events
 Morphogen gradients
 Soluble molecules which diffuse from source
cells and affect the differentiation of cells at a
distance
Conditional Cell Specification
Regeneration
 Both head and tail can be removed
from planaria
 Head and tail will regenerate from
anterior and posterior ends respectively
of trunk
 An anterior – posterior gradient allows
middle piece to “know” what parts to
regenerate on each end
 Conditional Cell
Specification
Regeneration in flatworms is
directed by an A-P morphogen
gradient
Conditional Cell Specification
Morphogen gradients
 Source – cell(s) that produce
morphogen
 Sink – site where morphogen is used
(degraded)
 Cells are either responding to varying
concentrations of morphogen or to
varying lengths of exposure times
 Conditional Cell Specification
Morphogen Gradients
& Thresholds

Cells in each flag differentiate

in response to the morphogen
gradient yet retain their original
specification as “French” or
“US”
 Conditional Cell Specification

Amphibian Morphogen Activin

Conditional Cell Specification
Conditional Cell Specification
Morphogenetic Fields
 Limb fields
 Region where cells are committed to form
a limb
 Cells within field are not determined to be
any particular part of the limb
 Other fields
 Eye & heart in vertebrates
 Imaginal discs in Drosophila
Conditional Cell Specification
Limb Field
Conditional Cell Specification
Invasion and splitting
of limb field by
nematode parasites
in frogs results in
duplication of limbs
Syncytial Cell Specification
 Syncytium
 Cytoplasm that contains multiple nuclei
 Many insect eggs begin development as
a syncytium – multiple nuclear divisions
occur before cellularization
 The syncytial cytoplasm contains
gradients of localized proteins and
mRNAs
 Upon cellularization, unequal amounts
of morphogens becomes localized
within cells
 Syncytial
Cell
Specification
Localized Maternal
mRNAs in Drosophila
establish protein
gradients in the
syncytial embryo
Syncytial Cell Specification
 The morphogens typically functioning in
syncytial specification are transcription
factors
 The unequal distribution of these factors
into cells of the embryo allow for the
specific activation or repression of
genes within the nuclei of the newly
formed cells
Cell Specification
 No embryo uses only one of the types
specification
 Regulative embryos use autonomous
specification frequently later in development
 Embryos that use autonomous specification
early use conditional specification for the
formation of some organs
 Syncytial specification can only be used for
very early events, both autonomous and
conditional specification occur later