Imaginal discs

The development of imaginal discs is a featureunique to the holometabolous insects

(beetles, moths, flies, and butterflies).
The life cycles of holometabolous insects include four stages: embryo, larva, pupa, and
adult (also known as imago).
Development through these stages does not occur simply by increasing the sizes of
exiting structures; instead, it involves a dramatic transformation from larva to adult.
The transformation is achieved by the replacement of larval tissues with a special
population of cells, known as imaginal discs, which form sac-like structures residing
inside the larva.
Imaginal discs have been studied most extensively in the fruit fly Drosophila
melanogaster
An imaginal disc is a sac-like epithelial structure found inside the larva of
insects that undergo metamorphosis. Once the larva turns into a pupa, almost all
the larval tissues degenerate and the imaginal discs turn into the external
structures of the head, thorax, limbs and genitalia.

The rest of the adult external structures (abdomen) originate from nests of
histoblast cells that are also formed in the larva.

The reason imaginal discs are of such interest to developmental biologists is that
they provide a powerful system to study multiple aspects of development and
biological mechanisms underlying disease; they are one of the reasons that make
the fruit fly Drosophila melanogaster an excellent model organism.
• Studies in Drosophila have shown that imaginal discs are initially set aside during embryogenesis,
apart from most embryonic cells that will contribute to the larva.

• In the embryo, the imaginal disc precursor cells can be visualized by the expression of molecular
markers or by their particular sizes, shapes, and movement.

• The imaginal discs appear in the newly hatched larva as local thickenings of the epidermis and, in the
case of the wing disc, contain around 40 cells

• the imaginal disc cells proliferate rapidly during larval stages. By the end of the larval stage, the largest
imaginal disc, that of the wing, contains approximately 50,000 cells

• During pupation, the imaginal disc cells go through morphogenetic movements, in response to a pulse
of the molting hormone ecdysone, to give rise to specific adult structures, accompanied by the
breaking-down of the larval tissues.
There are 19 discs in total: the epidermis of the head, thorax and limbs of the fly
come from 9 bilateral pairs of discs and the genitalia come from a medial disc.

Their names are: labial, clypeolabral, eye-antenna (these three will form the head);
humeral, wing, haltere (these will

form the notum, wings and halteres);

1st leg, 2nd leg, 3rd leg and

the genital disc

Drosophila imaginal discs regenerate after fragmentation.

(a) Imaginal discs resemble flattened sacs and are composed of two epithelial layers.
The disc proper has columnar cells, and the peripodial epithelium has squamous cells.
The apical surfaces of both layers point toward the lumen of the sac.

(c) Following fragmentation of the wing imaginal disc into a large three-quarters
fragment and a small one-quarter fragment, the large fragment regenerates the missing
portion, whereas the smaller fragment generates a mirror image of itself.

(d) In the case of the first leg (the foreleg), the smaller fragment from the anterior dorsal
quadrant regenerates the remainder of the disc, whereas the much larger three-quarters
fragment duplicates itself.
The pairs of eye-antenna, wings, and halteres come together, and the latter two
also fuse with the three pairs of leg discs. It is at this moment that they form a
continuous epithelium that has the recognisable shape of the adult fly.

An imaginal disc is a two-layered structure formed by a continuous epithelium.

One side is formed by the peripodial epithelium, a layer of flat and wide
squamous cells; the other side is a columnar epithelium comprising tall and thin
cells and designated as the disc proper.
The study of regeneration in Drosophila imaginal discs provides an opportunity to use
powerful genetic tools to address fundamental problems pertaining to tissue regeneration
and cell plasticity.
Discs respond to tissue damage in several ways:
(a) Removal of part of the disc elicits localized cell proliferation and regeneration of the
missing tissue.
(b) Damage at specific locations in the disc can cause cells to generate disc-
inappropriate structures (e.g., wing instead of leg), a phenomenon known as
transdetermination.
(c) Diffuse damage to imaginal discs, results in compensatory proliferation of surviving
cells.
Jun N-terminal kinase (JNK) signaling regulates multiple aspects of
regeneration.

Although high levels of JNK signaling have been shown to induce apoptosis,
JNK signaling is involved in wound healing during early stages of tissue
regeneration, which includes the formation of actin cables and filopodia.

JNK promotes tissue remodeling and cell proliferation.

JNK signaling likely increases cellular plasticity during regeneration.

JNK signaling also promotes a pupal delay

The fate of imaginal disc precursor cells is determined by intersecting anterior-
posterior and dorsal-ventral signals in the embryo.

These signals are provided by two sets of patterning genes along the embryonic
anterior-posterior and dorsal-ventral axes. Along the anterior-posterior axis, a
regulatory hierarchy involving gap genes, pair-rule genes, and segment polarity genes
determine the different cell fates.

The activities of several segment polarity genes, including wingless (wg), engrailed
(en), patched (ptc), and hedgehog (hh), are required for the consolidation of the
parasegment boundaries, which appear during early development of the embryo
imaginal discs increase their number of cells by thousands of fold during larval development.

Misregulation of disc growth can result in imaginal discs with abnormal sizes and
morphology.

Genetic studies have identified a large number of mutants that are either defective or
excessive in disc growth.

The disc overgrowth mutants are of particular interest, because they may provide genetic
models for tumor suppressor genes in humans.

Recessive mutations in several dozens of genes have been identified that cause two types of
disc overgrowth phenotypes: hyperplasia and neoplasia.

Hyperplastic discs retain the normal epithelial organization and the capacity to differentiate,
whereas the neoplastic discs do not.
imaginal discs are subdivided into distinct cell populations, called compartments.

• The compartmental boundary serves as a line of lineage restriction, and cells in

one compartment do not intermingle with cells in the other.

• The anterior and posterior compartments of the thoracic discs can be traced
back to the embryonic stage when the disc precursor cells are specified initially
along the anterior-posterior axis of the embryos

• Although patterning of all imaginal discs share some common features, the
identity of an individual imaginal disc is specified by the homeotic and other
selector genes.
• Studies of imaginal disc development in Drosophila have shed new light on
various aspects of vertebrate development. For example, gene expression and
functional studies suggest a striking similarity between fly appendages and
vertebrate limbs

• Several key regulatory genes in patterning imaginal discs have been implicated
in human diseases. For example, mutations in the human patched gene are
responsible for basal cell carcinoma, a common form of human cancer