Cell Fate

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The fate of a cell describes its future identity, or the identity of its daughter cells,

before it is actually phenotypically detectable through differentiation or division. Cell


fate can be determined by endogenous developmental factors, interaction with
adjacent cells, or external long-distance signals such as morphogens and hormones

The determination of a cell to a particular fate can be broken down into two states where the cell can
be specified (committed) or determined. In the state of being committed or specified, the cell type
is not yet determined and any bias the cell has toward a certain fate can be reversed or transformed
to another fate. If a cell is in a determined state, the cell's fate cannot be reversed or transformed. In
general, this means that a cell determined to differentiate into a brain cell cannot be transformed
into a skin cell. Determination is followed by differentiation, the actual changes in biochemistry,
structure, and function that result in specific cell types. Differentiation often involves a change in
appearance as well as function

Modes of specification[edit]
There are three general ways a cell can become specified for a particular fate; they are
autonomous specification, conditional specification and syncytial specification.[16]

Autonomous specification[edit]
This type of specification results from cell-intrinsic properties; it gives rise to mosaic development.
The cell-intrinsic properties arise from a cleavage of a cell with asymmetrically expressed maternal
cytoplasmic determinants (proteins, small regulatory RNAs and mRNA). Thus, the fate of the cell
depends on factors secreted into its cytoplasm during cleavage. Autonomous specification was
demonstrated in 1887 by a French medical student, Laurent Chabry, working on tunicate embryos. [17]
[18]
This asymmetric cell division usually occurs early in embryogenesis.

Conditional specification[edit]
In contrast to the autonomous specification, this type of specification is a cell-extrinsic process that
relies on cues and interactions between cells or from concentration-gradients of morphogens.
Inductive interactions between neighboring cells is the most common mode of tissue patterning. In
this mechanism, one or two cells from a group of cells with the same developmental potential are
exposed to a signal (morphogen) from outside the group

Syncytial specification[edit]
Main article: Syncytium
This type of a specification is a hybrid of the autonomous and conditional that occurs in insects. This
method involves the action of morphogen gradients within the syncytium. As there are no cell
boundaries in the syncytium, these morphogens can influence nuclei in a concentration-dependent
manner.
he Arabidopsis root epidermis is composed of two cell types whose pattern of
differentiation is directed by positional cues during development. Examination of
mutations has identified genes involved in the establishment of cell fate specification
in this tissue. TRANSPARENT TESTA GLABRA (TTG) and GLABRA2 (GL2) are
positive regulators of non-hair fate and are active during the early differentiation of
the epidermis in the meristem. GL2 encodes a homeobox protein which is
expressed in non-hair cells in the meristem and is positively regulated by TTG.
Mutations in genes involved in the regulation of ethylene biosynthesis and signal
transduction indicate that ethylene is a positive regulator of hair cell fate. Treatment
of ttg and gl2 plants with modulators of ethylene biosynthesis indicate that ethylene
acts down stream of TTG and GL2 during the fate specification process. The
relationship between meristem organisation and the mechanism underpinning the
establishment of cell fate in other systems is also discussed.

Cell lineage denotes the developmental history of a tissue or organ from the fertilized embryo. [1] Cell
lineage is based on the tracking of an organisms cellular ancestry due to the cell divisions and
relocation as time progresses, this starts with the originator cells and finishing with a mature cell that
can no longer divide. [2]Cell lineage can be studied by marking a cell (with fluorescent molecules or
other traceable markers) and following its progeny after cell division.

History of cell lineage[edit]


One of the first studies of cell lineages took place in the 1870s by Whitman who studied cleavage
patterns in leeches and small invertebrates.[6] He found that some groups, such as nematode worms
and ascidians form a pattern of cell division which is identical between individuals and invariable.
This high correlation between cell lineage and cell fate was thought to be determined by segregating
factors within the dividing cells. Other organisms had stereotyped patterns of cell division and
produced sublineages which were the progeny of particular precursor cells. These more variable cell
fates are thought to be due to the cells' interaction with the environment. [6]
Techniques of fate mapping[edit]
Further information: Fate mapping
Cell lineage can be determined by two methods, either through direct observation or through clonal
analysis. During the early 19th century direct observation was used however it was highly limiting as
only small transparent samples could be studied. With the invention of the confocal microscope this
allowed larger more complicated organisms to be studied. [6]
Perhaps the most popular method of cell fate mapping in the genetic era is through site-specific
recombination mediated by the Cre-Lox or FLP-FRT systems. By utilizing the Cre-Lox or FLP-FRT
recombination systems, a reporter gene (usually encoding a fluorescent protein) is activated and
permanently labels the cell of interest and its offspring cells, thus the name cell lineage tracing [7].
With the system, researchers could investigate the function of their favorite gene in determining cell
fate by designing a genetic model where within a cell one recombination event is designed for
manipulating the gene of interest and the other recombination event is designed for activating a
reporter gene. One minor issue is that the two recombination events may not occur simultaneously
thus the results need to be interpreted with caution[8]. Furthermore, some fluorescent reporters have
such an extremely low recombination threshold that they may label cell populations at undesired
time-points in the absence of induction [9].

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