Insects[edit]

Further information: Insect anatomy § Antennae

Terms used to describe shapes of insect antennae

Antennal shape in the Lepidoptera from C. T. Bingham (1905)

Insects evolved from prehistoric crustaceans, and they have secondary antennae like crustaceans,
but not primary antennae. Antennae are the primary olfactory sensors of insects[6] and are
accordingly well-equipped with a wide variety of sensilla (singular: sensillum). Paired, mobile, and
segmented, they are located between the eyes on the forehead. Embryologically, they represent the
appendages of the second head segment.[7]
All insects have antennae, however they may be greatly reduced in the larval forms. Amongst the
non-insect classes of the Hexapoda, both Collembola and Diplura have antenna, but Protura do not.
[8]

Antennal fibrillae play an important role in Culex pipiens mating practices. The erection of these
fibrillae is considered to be the first stage in reproduction. These fibrillae serve different functions
across the sexes. As antennal fibrillae are used by female C. pipiens to locate hosts to feed on,
male C. pipiens utilize them to locate female mates.

Structure[edit]
Electron micrograph of antenna surface detail of a wasp (Vespula vulgaris)

The three basic segments of the typical insect antenna are the scape or scapus (base),
the pedicel or pedicellus (stem), and finally the flagellum, which often comprises many units known
as flagellomeres.[9] The pedicel (the second segment) contains the Johnston's organ which is a
collection of sensory cells.
The scape is mounted in a socket in a more or less ring-shaped sclerotised region called
the torulus, often a raised portion of the insect's head capsule. The socket is closed off by the
membrane into which the base of the scape is set. However, the antenna does not hang free on the
membrane, but pivots on a rigidly sprung projection from the rim of the torulus. That projection on
which the antenna pivots is called the antennifer. The whole structure enables the insect to move
the antenna as a whole by applying internal muscles connected to the scape. The pedicel is flexibly
connected to the distal end of the scape and its movements in turn can be controlled by muscular
connections between the scape and pedicel. The number of flagellomeres can vary greatly between
insect species, and often is of diagnostic importance.
True flagellomeres are connected by membranous linkage that permits movement, though the
flagellum of "true" insects does not have any intrinsic muscles. Some other Arthropoda do however
have intrinsic muscles throughout the flagellum.
Such groups include the Symphyla, Collembola and Diplura. In many true insects, especially the
more primitive groups such as Thysanura and Blattodea, the flagellum partly or entirely consists of a
flexibly connected string of small ring-shaped annuli. The annuli are not true flagellomeres, and in a
given insect species the number of annuli generally is not as consistent as the number of
flagellomeres in most species.[9]
In many beetles and in the chalcidoid wasps, the apical flagellomeres form a club shape, and the
collective term for the segments between the club and the antennal base is the funicle; traditionally
in describing beetle anatomy, the term "funicle" refers to the segments between the club and
the scape. However, traditionally in working on wasps the funicle is taken to comprise the segments
between the club and the pedicel.[9]
Quite commonly the funicle beyond the pedicel is quite complex in Endopterygota such as beetles,
moths and Hymenoptera, and one common adaptation is the ability to fold the antenna in the middle,
at the joint between the pedicel and the flagellum. This gives an effect like a "knee bend", and such
an antenna is said to be geniculate. Geniculate antennae are common in the Coleoptera and
Hymenoptera. They are important for insects like ants that follow scent trails, for bees and wasps
that need to "sniff" the flowers that they visit, and for beetles such
as Scarabaeidae and Curculionidae that need to fold their antennae away when they self-
protectively fold up all their limbs in defensive attitudes.
Because the funicle is without intrinsic muscles, it generally must move as a unit, in spite of being
articulated. However, some funicles are complex and very mobile. For example, the Scarabaeidae
have lamellate antennae that can be folded tightly for safety or spread openly for detecting odours
or pheromones. The insect manages such actions by changes in blood pressure, by which it exploits
elasticity in walls and membranes in the funicles, which are in effect erectile. [10]
In the groups with more uniform antennae (for example: millipedes), all segments are
called antennomeres. Some groups have a simple or variously modified apical or subapical bristle
called an arista (this may be especially well-developed in various Diptera).[11]

Functions[edit]
Olfactory receptors (scales and holes) on the antenna of the butterfly Aglais io, electron micrograph

Olfactory receptors on the antennae bind to free-floating molecules, such as water vapour,
and odours including pheromones. The neurons that possess these receptors signal this binding by
sending action potentials down their axons to the antennal lobe in the brain. From there, neurons in
the antennal lobes connect to mushroom bodies that identify the odour. The sum of the electrical
potentials of the antennae to a given odour can be measured using an electroantennogram.[12]
In the monarch butterfly, antennae are necessary for proper time-compensated solar
compass orientation during migration. Antennal clocks exist in monarchs, and they are likely to
provide the primary timing mechanism for sun compass orientation.[13]
In the African cotton leafworm, antennae have an important function in signaling courtship.
Specifically, antennae are required for males to answer the female mating call. Although females do
not require antennae for mating, a mating that resulted from a female without antennae was
abnormal.[14]
In the diamondback moth, antennae serve to gather information about a host plant's taste and odor.
After the desired taste and odor has been identified, the female moth will deposit her eggs onto the
plant.[15] Giant swallowtail butterflies also rely on antenna sensitivity to volatile compounds to identify
host plants. It was found that females are actually more responsive with their antenna sensing, most
likely because they are responsible for oviposition on the correct plant. [16]
In the crepuscular hawk moth (Manduca sexta), antennae aid in flight stabilization. Similar
to halteres in Dipteran insects, the antennae transmit coriolis forces through the Johnston's organ
that can then be used for corrective behavior. A series of low-light, flight stability studies in which
moths with flagellae amputated near the pedicel showed significantly decreased flight stability over
those with intact antennae.[17] To determine whether there may be other antennal sensory inputs, a
second group of moths had their antennae amputated and then re-attached, before being tested in
the same stability study. These moths showed slightly decreased performance from intact moths,
indicating there are possibly other sensory inputs used in flight stabilization. Re-amputation of the
antennae caused a drastic decrease in flight stability to match that of the first amputated group.