Antennal lobe neurons
Neurons in the antennal lobe are primarily olfactory receptor
neurons (their axons), local interneurons whose neurites are
entirely confined to the antennal lobe, and antennal lobe
projection neurons, which represent the output of the antennal
lobe. A small number of efferent processes of neurons from other
brain areas are also present.
Olfactory receptor
neurons
Olfactory receptor neurons projecting to the
antennal lobe are originating from the antennae in most insects and
also from the maxillary palp (a moth part) in some more derived
groups such as Lepidoptera and Diptera. Their somata are located in
proximity to the sensillar structures. Olfactory receptor neurons
expressing the same odorant receptor project to the same glomeruli
[Vosshall et al., 2000].Generally,
a large number of olfactory recpetor neurons converge onto a
projection neuron and synapses are located in the outer “cap”
of the glomerulus.
Projection neurons
Projection
neurons have dendrites in the antennal lobe and axons projecting to
the protocerebrum. There are projection neurons whose dendrites are
confined to a single glomerulus (uniglomerular) and projection
neurons whose dendrites innervate more than one glomerulus
(multiglomerular). In general, there are three projection tracts to
the protocerebrum called antenno-cerebral tracts (ACTs) [Galizia &
Rössler, 2010].In
the tobacco hornworm moth, a detailed analysis of these tracts has
been made [Homberg et al., 1988]. In the silkmoth, it has also been
shown that there is a topographic relationship between glomeruli,
ACTs and projection areas in the protocerebrum [Kanzaki et al.,
2003].In
most insects, uniglomerular projection neurons are the predominant
type. In the locust (which has an unusually large number of
glomeruli), multiglomerular projection neurons are typical.
Local
interneurons
The neurites of antennal lobe interneurons are
entirely confined to the antennal lobe. These cells are considered
to be the main element of information processing in the antennal
lobe. In the silkmoth, the first extensive morphological survey of
local interneurons was provided. In combination with
immunohistochemistry for GABA (an inhibitory neurotransmitter), it
could be shown that local interneurons innervating many glomeruli
are inhibitory. The study also showed a diversity of types of local
interneurons [Seki & Kanzaki, 2008].In
the honey bee, a social insect, the morphology of local
interneurons appears to be more complex.
Efferent
innervation
Efferent innervation of the antennal lobe is
less well studied that the other neuronal constituents. This is
presumably due to the fact that the number of cells efferently
innervating the antennal lobe is small.
In the honey bee,
some putatively octopaminergic ventral unpaired median neurons
whose somata are located in the suboesophageal ganglion, innervate
the antennal lobe. For one of them, it has been shown that its
activation can replace the unconditioned stimulus in classical
olfactory conditioning [Hammer , 1997].
Efferent innervation by serotonergic neurons is known for a number of insect
species [Dacks et al., 2006], including the silkmoth [Hill et al., 2002](serotonin immunoreactive antennal lobe feedback neuron in silkmoth).
References
Dacks
AM, Christensen TA, Hildebrand JG (2006) Phylogeny of a
serotonin-immunoreactive neuron in the primary olfactory center of
the insect brain. J Comp Neurol 498:727-746.
Galizia CG,
Rössler W (2010) Parallel olfactory systems in insects: anatomy
and function. Annu Rev Entomol 55:399-420.
Hammer M (1997)
The neural basis of associative reward learning in honeybees.
Trends Neurosci 20:245-252.
Hill ES, Iwano M, Gatellier L and
Kanzaki R (2002) Morphology and physiology of the
serotone-immunoreactive putative antennal lobe feedback neuron in
the male silkmoth Bombyx mori. Chemical Senses 27:475-483.
Homberg
U, Montague RA, Hildebrand JG (1988) antenna-cerebral pathways in
the brain of the sphinx moth Manduca sexta. Cell Tissue Res
254:255-281.
Seki Y, Kanzaki R (2008) Comprehensive
morphological identification and GABA immunocytochemistry of
antennal lobe local interneurons in Bombyx mori. J Comp Neurol 506:
93-107.
Vosshall LB, Wong AM, Axel R (2000) An olfactory
sensory map in the fly brain. Cell 102:147159.