Types of sensilla
Two major types of sensilla can be distinguished in the fully developed
embryo - external sensory organs, located within the epidermis, and chordotonal
organs, located internally in the body wall or close to the somatic muscles.
External sensilla contain one or more bipolar neurons, enclosed by three
accessory cells. The cell bodies of bipolar neurons lie beneath the epidermis;
their dendrite(s) penetrate the epidermis and terminate in conjunction
with the cuticle. Accessory cells are modified epidermal cells which form
concentric sheaths around the dendrites and build the stimulus receptor
structure. The outer accessory cells form the socket (tormogen cell) and
the shaft (trichogen cell) of the sensory organ. The inner cell (thecogen
cell) secretes a cuticle-like matrix around the tip of the dendrite(s),
called the dendritic cap. Most external sensilla are mechanoreceptors;
these include hairs (trichoid sensilla) and an ill-defined type called
a "papilla sensillum" or "campaniform sensillum. Mechanoreceptors typically
possess a single sensory neuron, although one hair and one papilla sensillum
of each abdominal segment are innervated by three neurons each. Another
type of sensillum which occurs only in the thorax and the telson has been
called a basiconical sensillum, characterized by a club shaped process
in an epidermal depression. Basiconical sensilla are innervated by three
sensory neurons and are probably hygroreceptors.
Chordotonal organs are stretch receptors attached to the basal (inner)
surface of the epidermis. In the embryo, chordotonal organs are formed
by one, three or five identical units called scolopidia. Each scolopidium,
like the external sensillum, has one bipolar neuron and three accessory
cells. Unlike those of external sensilla, accessory cells of chordotonal
organs are not organized as sheaths around the sensory neurons, but form
ligamentous structures by means of which the chordotonal organs insert
at the epidermis.
In addition to the sensilla, which comprise bipolar neurons and accessory
cells, several different types of isolated sensory neurons, i.e., devoid
of accessory cells, are present internally in the embryo. The neurons
possess more than one dendrite, and are therefore referred to as multidendritic
neurons. Multidendritic neurons occur as clusters of up to five cells
attached to the basal surface of the epidermis or internal organs, such
as tracheae, peripheral nerves, and muscles. Three types of multidendritic
neurons can be distinguished: those with arborizations underneath the
epidermis (da neurons), those innervating the tracheal branches
(td neurons), and those with two opposing dendrites (bd neurons).
As a rule, neurons and accessory cells of a given sensillum are derivatives
of the same sensory organ progenitor cell (SOP). The progenitor cells
of external organs innervated by a single neuron undergo two divisions.
The progenitors of chordotonal organs and multidendritic neurons delaminate
from the epidermal anlage into the interior of the embryo. Proliferation
of the SOPs of chordotonal organs and multidendritic neurons begins after
they have delaminated and become attached to the basal surface of the
ectoderm. Following delamination, the progenitors of multidendritic neurons
undergo one to three further divisions to form clusters of up to five
neurons. Progenitors of the tri- and pentascolopidial chordotonal organs
have a highly complex proliferation pattern which has not been fully reconstructed.
The pattern of proliferation of the progenitors of multiply innervated
external sensilla is controversial.
Small head sensilla
The dorsal pharyngeal organ (dpo) and a dorsal
pharyngeal monoscolopidial chordotonal organ (pch1) are both derived
from the labral segment. The dpo is located posterior to the labral
sensory complex and is innervated by three neurons of unknown sensory
modality. Two sensory structures, the papilla organ (pao) and an
associated organ (ao) are spatially related to the terminal organ.
The pao is probably of mandibular origin. It comprises a multiply
innervated sensillum in the lateral atrial wall. The hypomaxillary organ
(hmo) is a multiply innervated sensillum of unknown sensory modality
and segmental origin. It is located close to the labial organ, but its
axon projects with the maxillary nerve. The hypopharyngeal organ and the
latero-hypopharyngeal organ (lho) are likely derived from the intercalary
segment. Since these sensilla are among the first ones to differentiate
in the head, their movement from a position flanking the stomodeum (position
of the intercalary segment) to the floor of the pharynx can be easily
followed. The hypopharyngeal organ is innervated by three or four dendrites.
Its axons join an anteriorly directed bundle which runs to the labral
nerve; the two or three neurons of the latero-hypopharyngeal organ are
located at the junction of both branches.The lateropharyngeal organ (lpo)
appears at a relatively early stage in the dorsal ridge which is derived
by fusion from the dorsal parts of the labium and maxilla. During head
involution, the lpo becomes incorporated into the dorsal pouch.
The lpo is located at the lumen of the dorsal pouch, and consists
of two or three neurons.
Abdominal segments a1-a7
Ventral group: The ventral group consists of six papilla sensilla
(vp1, vp2, vp3, vp4, vp4a, vp5), two monoscolopidial chordotonal
organs (vch1 and vch2), and several clusters of multidendritic
neurons. The ventralmost papilla sensilla vp1 and vp2, are
located in the paraneural space at the posterior border of the corresponding
segmental nerve. The neurons of vp3, vch1 and vch2,
and vdap form a dense cluster located posteriorly in each segment
at the level of the ventral oblique musculature. The axons of vp1-3,
vch1 and vch2, and vdap, together with the motor
axons innervating the ventral oblique muscle fibres, form the branch "c"
of the segmental nerve. A side branch of this axon bundle carries sensory
axons of the multidendritic neurons vdaA-D and vbp. The
cells of vp4, vp4a, and vdaa are located in close
proximity to the intersegmental nerve within the anterior half of each
segment. Their axons join branch "b" of the segmental nerve which
also carries motor axons to several ventral longitudinal and oblique muscle
fibres. Finally, there is the group of three sensory neurons of vp5
and the multidendritic vdap neuron, which are the lateralmost representatives
of the ventral group of sensilla and which are located at the lateral
boundary of the ventral musculature. Their axons join branch "a"
of the segmental nerve.Two multidendritic neurons, the intersegmental
bidendritic neuron (isbd) and trachea-associated neuron (istd), are located
on the segment boundary. Axons of both neurons form a separate nerve,
called the transverse nerve which extends ventrally along the segment
boundary and enters the ventral nerve cord at a dorsomedial position.
Lateral group: The most prominent sensillum of the lateral group of
abdominal sensilla is lch5, the pentascolopidial chordotonal organ.
Its five scolopidia are located posteriorly in the niche formed between
the epidermis and the lateral transverse muscle. These scolopidia are
strictly aligned and oriented dorsoposteriorly. The ligament cells anchoring
lch5 to the epidermis are very long; they insert at close to the
posterior segment boundary at the level of the dorsal muscle fibres DO2.
During development the lch5 cluster is born in the dorsal ectoderm.
In the thoracic segments the lch5 cluster remains at this position
whereas the lch5 cluster moves ventrally in the abdominal segments.
Sensory neurons of three external sensilla, lh1, lh2 and
lp1, and two multidendritic neurons, ltd and lda,
flank the pentascolopidial chordotonal organ at its anterior border. The
monoscolopidial chordotonal organ lch1 is located more superficially
and dorsally than lch5, being oriented dorso-anteriorly. The axons
of all lateral sensilla, except for lch1, join the intersegmental
nerve; the lch1 axon projects to join branch "a" of the segmental
nerve.
Dorsal group: The dorsal group consists of four external sensilla,
a small and a large hair sensillum (dh1 and dh2) and two
papillar sensilla (dp1 and dp2), arranged along a vertical
line in the middle of each abdominal segment, between the epidermis and
the dorsal acute muscles. The small dorsal hair dh1 is innervated
by three neurons, the large dh2, as well as dp1 and dp2,
are each in contact with only one neuron. Sensory neurons of all four
sensilla form a dense cluster with five multidendritic neurons (ddaA-E).
A sixth multidendritic, bipolar neuron located ventral to this cluster
(dbp) forms conspicuous longitudinal processes which span the entire
segment. Axons of all dorsal sensilla course through the intersegmental
nerve.
Thoracic segments
Ventral group: The ventral group consists of five sensilla in each
thoracic segment: two papillar sensilla (vp1 and vp2), a
monoscolopidial chordotonal organ (vch1), a basiconical sensillum
(vbd) and Keilin's organ (ko). The cell bodies of the neurons
of vbd and vp2, together with four multidendritic neurons
(vdaA-D), are grouped in a somewhat irregularly shaped cluster
in the anterior half of the segment, at the level of the ventral oblique
muscles (VO1-3). The five neurons associated with Keilin's organ and the
monoscolopidial organ vch1 can be found more posteriorly. These
neurons form part of the leg imaginal disc and persist into the adult
period.
Lateral group: The patterns of sensilla of the lateral and dorsal
group differ in the three thoracic segments. In the prothorax, the lateral
group comprises a tight cluster of cells formed by papillar sensilla lp1
and lp2, hair sensillum lh1, a triscolopidial chordotonal
organ lch3, and four multidendritic neurons (ldaA-D). Strikingly,
the axons of lch3 join the segmental nerve of the mesothoracic
segment to course towards the CNS. In addition, there are two more papillar
sensilla (px, py) close to the anterior boundary of the
prothorax. In the mesothorax and metathorax these two papillae are absent
and instead of the lateral triscolopidial chordotonal organ, there is
a basiconical sensillum (lbd).
Dorsal group: In the prothorax, the dorsal group of sensilla comprises
hair sensilla dh1 and dh2, with one neuron each; three papillae,
dp1-dp3, with one neuron each; a triscolopidial chordotonal organ
(dch3), and a dorsal basiconical sensillum (dbd). Clustered
around the neurons of these sensilla, there are six multidendritic neurons
(ddaA-E; dbp). In mesothorax and metathorax, the dorsal basiconical
sensillum is missing.
The innervation of thoracic sensory organs follows the same scheme as
that of abdominal sensilla: the ventral group sends its axons through
two different branches into the posterior fascicle or the segmental nerve.
The axons of the lateral and dorsal group course through the intersegmental
nerve.
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