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Imaginal Discs: Quick Guide
Imaginal Discs: Quick Guide
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Quick guide
Imaginal discs
Silvia Aldaz1 and
Luis M. Escudero1,2
A Mouth part
B A P
Frontal plate and upper lip
A P
Antenna
Eye Head Wing
Prothorax
Leg
Thorax
Haltere Hinge
Abdomen
Wing
Notum
Genitalia
How do they develop into the adult epithelium that has the recognisable of all these factors forms a
epidermis? The changes needed shape of the adult fly. During ‘prepattern’, which presages the
to transform a small nest of cells metamorphosis, the imaginal cells final differentiated structure. Each
into a differentiated adult tissue are will continue to develop as different region of the disc will have a different
enormous and encompass most cell types, including, for example, combination of prepattern genes,
aspects of developmental biology mechanosensory organs, epidermal and these will determine different
(which is why imaginal discs are cells, wing veins or photoreceptors. cell fates. All this makes it possible
such a good model system). First to draw a ‘fate map’ in the discs
of all, the imaginal cells have a How do adult structures appear establishing a correlation between
high rate of proliferation. They can in stereotypical positions? the different disc regions and the
divide up to every 10 hours during This is the essence of developmental resulting adult structures (Figure 1B).
the 4 days of larval life. In the case biology! The developing imaginal
of the mature wing disc (the largest disc epithelium contains precise What is the three-dimensional
of all the discs), this results in a positional information, giving structure of the imaginal discs?
final size of approximately 50,000 rise to subdivisions of the disc The establishment of cell fate,
cells. Of course, this growth has to into distinct domains. The initial as outlined above, provides a
be carefully regulated. At the same subdivisions are the so-called rather misleading two-dimensional
time that these cells are growing, ‘compartments’ (anterior and description of disc development;
the different adult cell types are posterior, and ventral and dorsal). a canvas where the combination
determined. These are defined by cell-lineage of signals paint gene expression
Upon metamorphosis, the discs restrictions, meaning that cells patterns. In fact, imaginal discs
evert, simultaneously moving from born in one compartment cannot are bag-like structures, and they of
the inside of the larva through the cross into another, and they are course give rise to three- dimensional
larval wall to the free space created affected by the expression of specific organs. An imaginal disc is a
under the pupal case. The discs then transcription factors. The cells from two- layered structure formed by a
unfold and elongate, completing different compartments do not mix continuous epithelium (imagine an
the evagination, and continue the when the discs are growing. The empty hot water bottle), where the
final differentiation of the external next level of organization is that the two sides are distinct. One side is
structures of the adult. About 6 boundaries between compartments formed by the peripodial epithelium,
hours after pupariation, fusion form important ‘organizing centers’, a layer of flat and wide squamous
between bilateral pairs of discs becoming the source of long- and cells; the other side is a columnar
begins. The pairs of eye-antenna, short- range diffusible signals known epithelium comprising tall and thin
wings, and halteres come together, as morphogens. The morphogens cells and designated as the disc
and the latter two also fuse with the are able to control the expression proper. The cells at the junction
three pairs of leg discs. It is at this of transcriptional activators and between the two sheets have an
moment that they form a continuous repressors. The combination intermediate cuboidal morphology.
Magazine
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Extending the analogy of the hot Coriolis effect. The resulting irregular
water bottle, the inside of the bottle Primer convection cells and gyres are aligned
corresponds to the apical surfaces roughly north–south and generate
of the squamous and columnar most of the field felt by animals at
epithelia. Magnetoreception the earth’s surface. Because of the
Nearly all research into imaginal geometry of the field, the magnetic
disc development has been focused James L. Gould force lines at most locations slant
on the columnar cells of the disc down toward the nearer pole at an
proper — after all, they make almost Few subjects in animal behavior angle called the ‘dip’.
all of the adult organs. But it is have more exotic mystery than As might be expected of an
becoming clear that the peripodial magnetic- field sensitivity. A force we electromagnet created in such a
epithelium cannot be ignored: it cannot sense, generated by events haphazard way, the earth’s magnetic
is essential for the eversion and no one completely understands, field varies over time in average
fusion of the discs. There is also creates field lines that pass through strength, direction, and even
evidence that peripodial cells are our bodies without any evident effect polarity. Near the short-term end of
required for growth, patterning on us or on them. It is an energy felt the scale, the north magnetic pole
and morphogenesis of the disc as much by migrating lobsters on has wandered about 1.4° north and
proper. So we can begin to think the sea floor as by ocean- crossing 3.6° west just in the last five years;
of the two-layered disc as a more birds thousands of meters overhead, this corresponds to a migration of
complex tissue in its own right, with transduced in generally poorly about 175 m per day. Shifting cells
interactions between the layers understood ways. Despite the and gyres are presumably at fault.
driving development of the structures blindness of humans, modern life At the long-term extreme, complete
that will become the final adult depends on this invisible, ghostlike reversals occur (very) roughly every
organs. field. Aside from lights and heaters, 300,000 years, though the last was
nearly every electrical device we own 780,000 years ago; a reversal now
What is the future for imaginal makes use of electromagnetism, and would be extremely inconvenient
disc studies? Of course we can’t that same magnetism is essential in to humans and other animals alike.
know what will happen in the future. generating the power these new- found Clearly, at least some creatures
But recent advances make it clear necessities consume. But for many needing to make use of such a
that the live observation of the animals, the reliance is far older and fickle field must be able to calibrate
previously described processes is on more basic: their life-or- death ability themselves to the field strength
the horizon and this will make a big to find their way around in the world and direction at least once in their
difference. After all, the processes depends on correctly interpreting the lifetime; some do so daily [1].
that transform an undifferentiated earth’s magnetic field. While this dynamo theory of
bag of 50 cells into a complex circulating metals helps account for
three- dimensional adult organ are Magnetism more than 95% of the magnetism,
highly dynamic; the ability to study As any survivor of introductory there is a much more variable
them in real time in future, rather college physics knows, a flow of residual component with clear
than by snapshots of fixed tissue, electrons generates a magnetic short-term effects on at least homing
should have a dramatic effect on field around it, while a changing or pigeons and honey bees. The earth’s
the next few years of imaginal disc travelling magnetic field induces jet streams capture ions from the
research. movement in nearby electrons. Each ionosphere and move them eastward
electron is a tiny magnetic-field at up to hundreds of kilometers per
Where I can find out more? generator: it has a spin (of which hour. As moving electrons, they
Bates, M. and Martínez-Arias, A. (1993). The there are two quantum-mechanical generate their own magnetic field.
Development of Drosophila melanogaster.
(New York: Cold Spring Harbor Laboratory.) states, ‘up’ and ‘down’) plus, if it is The jet streams shift north and
García-Bellido, A., Ripoll, P. and Morata, G. part of an atom, an orbital motion south on a daily basis simply as a
(1973). Developmental compartmentalisation
of the wing disc of Drosophila. Nature New relative to its nucleus. The spinning consequence of the warming and
Biol. 245, 251–253. produces roughly twice the field cooling (and consequent expansion
Gibson, M.C., and Schubiger G. (2000). strength of orbiting. These two and contraction) of the atmosphere.
Peripodial cells regulate proliferation and
patterning of Drosophila imaginal discs. Cell fields are not normally directionally From day to day and week to week
103, 343–350. correlated. For most biomagnetic they can migrate across hundreds
Held, L.I. (2002). Imaginal Discs, the Genetic effects, spin is the key element. of kilometers of latitude, changing
and Cellular Logic of Pattern Formation.
(Cambridge: Cambridge University Press.) The precise origin of the earth’s not only the weather below but the
Hartenstein, V. (1993). Atlas of Drosophila. (New field is still controversial, though strength and direction of the small
York: Cold Spring Harbor Laboratory.)
the broad outlines seem clear (by magnetic field they induce. During
exclusion if for no better reason). magnetic ‘storms’ the sun delivers
1MRC Laboratory of Molecular Biology, The earth’s outer core consists huge numbers of new ions to the jet
Hills Road, Cambridge CB2 2QH, UK. of a flowing but viscous metallic streams, briefly and unpredictably
2Current address: Hospital Virgen Del Rocío/
liquid. The convective transfer of amplifying this induced field [2].
CSIC/Universidad de Sevilla.
heat from the radioactive inner
Instituto Biomedicina Sevilla. Av.
Manuel Siurot, Sevilla 41013, Spain. core along with the rotation of the Maps and compasses
E-mail: saldaz@mrc-lmb.cam.ac.uk; earth keeps the molten iron core in Aside from evidence that honey
luisma.escudero@googlemail.com motion, probably in part through the bees can use the diurnal variations