Detection of apoptosis
DNA cleavage has been one of the main biochemical criteria used to
distinguish apoptosis from necrosis.However,it should not be the sole
factor used.Both morphological and biochemical characteristics need to be
exemined collectively to determine that cell death hasoccured via
apoptosis.The main methods methods of apoptosis detection are summarized
below.
Microscopy
Histological sections can be treated with reagents(e.g.haematoxylin and
eosin)that have an affinity for negatively charged molecules and stain the
nuclear material,revealing its distribution.When there sections are examined
under a light microscope,apoptosis is shown by shrinkage of the nuclei of
the cells and apoptotic bodies show up as small darkly stained structures.
By counting apoptic cells before and a given treatment,it is possible
to asses the efficacy of that treatment in inducing apoptosis and how readily
a given tissue or tumour undergoes the process.Alternatively, cells grown
in culture can be examined by scanning microscopy,which shows changes in the
shape of individual cells undergoing apoptosis and the characteristic
blebing on the cell's surface.
Flow cytometry
Cells in suspension are stained with fluorescent antibodies by attachment
of these antibodies to the cell surface antigens.A laser beam is directed
at the cell,causing the antibody to fluoresce.The intensity of the
fluorescense reveals the number of cells and the amount of scattering of
the beam depends on the size of the cells.When cells undergo apoptosis
they shrink and break into apoptotic bodies.This is revealed as a reduction
in light scattering by the cell sample.
DNA Electrophoresis
DNA is isolated from cells,placed on a gel and an electric current is
passed through the gel.Since DNA has a negative charge.the electric
current will cause the DNA to migrate depending on its net charge,size
and shape. The DNA is placed on a solution with a powerful negatively
charged detergent.The DNA then migrates towards a positive electrode when
a voltage is applied.The different fragments of DNA migrate across the
gel according to their size,with small fragments moving much more readily
than large ones.The result is a series of descrete bands arranged in
order of molecular weight,which can be detected by photographing the gel
under ultraviolet light.The sizes of the fragments of DNA are calculatde
with refrence to markers of known size run along the sample.
Apoptosis is revealed by the
breakdown of the DNA into fragments.This hallmark of apoptosis proceeds
in two steps:
- First there is relatively infrequent cutting_producing fragments in the
range 300bp-50kbp.
- Then the DNA cleavage continues-resulting in a
characteristic"fragmentation ladder"ofDNA bands seen on the gel.These
bands represent DNA of various molecular weights which are multiples of
180bp.These fragments correspond to nucleosomes,therefore this process is
described as internucleosomal fragmentation.
In vitro experiments suggest that the initial breakage of DNA into large
fragments is of crucial importance to apoptosis,whereas the DNA cleavage
into smaller fragments now appears to be a by-product of the apoptotic
process.These experiments have demonstrated that cells undergo the
morphological changes characteristic of apoptosis despite selective
inhibition of internucleosomal DNA fragmentation by intracellular Ca+2
chelation or by treatment with phorbol ester.However,in contrast,neither
apoptopic morphology more internucleosomal DNA fragmentation was blocked
by inhibition of nucleoside analogue incorporation into DNA.(Huang et
al.,in press).
Detection of apoptosis in patients
Most work on apoptosis has been done in animals,cell samples or cell
cultures.If apoptosis could be detected in tissue samples from cancer
patients,this would support the clinical significance of this process,
particularly if a treatment colud be shown to induce apoptosis in tumor
cells.To date it has not been possible to isolate apoptotic cells in blood
or tissue taken from patients after treatment in vivo,probably because
apoptotic cells are rapidly ingested and destroyed by phagocytes and
neutrophils.Recently,however,large nuclesomal fragmentation has been
detected in leukaemic lymphocytes from patients after chemotheraphy fpr CLL
(Huang et al.,submitted).