Insect mouthparts
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Insects have
mouthparts Mouthparts may refer to: * The parts of a mouth ** Arthropod mouthparts The mouthparts of arthropods have evolved into a number of forms, each adapted to a different style or mode of feeding. Most mouthparts represent modified, paired append ...
that may vary greatly across insect species, as they are adapted to particular modes of feeding. The earliest insects had chewing mouthparts. Most specialisation of mouthparts are for piercing and sucking, and this mode of feeding has evolved a number of times idependently. For example, mosquitoes and aphids (which are
true bugs Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from to aroun ...
) both pierce and suck, however female mosquitoes feed on animal blood whereas aphids feed on plant fluids.


Evolution

Like most external features of arthropods, the mouthparts of
Hexapoda The subphylum Hexapoda (from Greek for 'six legs') comprises most species of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura (all of these were once considere ...
are highly derived. Insect mouthparts show a multitude of different functional mechanisms across the wide diversity of insect species. It is common for significant homology to be conserved, with matching structures forming from matching
primordia A primordium (; plural: primordia; synonym: anlage) in embryology, is an organ or tissue in its earliest recognizable stage of development. Cells of the primordium are called primordial cells. A primordium is the simplest set of cells capable o ...
, and having the same evolutionary origin. However, even if structures are almost physically and functionally identical, they may not be homologous; their analogous functions and appearance might be the product of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.


Chewing insects

Examples of chewing insects include dragonflies, grasshoppers and
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s. Some insects do not have chewing mouthparts as adults but chew solid food in their larval phase. The moths and butterflies are major examples of such adaptations.


Mandible

A chewing insect has a pair of mandibles, one on each side of the head. The mandibles are caudal to the labrum and anterior to the maxillae. Typically the mandibles are the largest and most robust mouthparts of a chewing insect, and it uses them to masticate (cut, tear, crush, chew) food items. Two sets of muscles move the mandibles in the
coronal plane The coronal plane (also known as the frontal plane) is an anatomical plane that divides the body into dorsal and ventral sections. It is perpendicular to the sagittal and transverse planes. Details The coronal plane is an example of a longitud ...
: abductor muscles move insects' mandibles apart ( laterally); adductor muscles bring them together ( medially). This they do mainly in opening and closing their jaws in feeding, but also in using the mandibles as tools, or possibly in fighting; note however, that this refers to the coronal plane of the mouth, not necessarily of the insect's body, because insects' heads differ greatly in their orientation. In carnivorous chewing insects, the mandibles commonly are particularly serrated and knife-like, and often with piercing points. In herbivorous chewing insects mandibles tend to be broader and flatter on their opposing faces, as for example in caterpillars. In males of some species, such as of Lucanidae and some Cerambycidae, the mandibles are modified to such an extent that they do not serve any feeding function, but are instead used to defend mating sites from other males. In some ants and
termite Termites are small insects that live in colonies and have distinct castes ( eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blat ...
s, the mandibles also serve a defensive function (particularly in soldier castes). In bull ants, the mandibles are elongate and toothed, used both as hunting and defensive appendages. In bees, that feed primarily by use of a proboscis, the primary use of the mandibles is to manipulate and shape wax, and many paper wasps have mandibles adapted to scraping and ingesting wood fibres.


Maxilla

Situated beneath (caudal to) the mandibles, paired maxillae manipulate and, in chewing insects, partly masticate, food. Each maxilla consists of two parts, the
proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position ...
cardo (plural ''cardines''), and distal stipes (plural ''stipites''). At the apex of each stipes are two lobes, the inner lacinia and outer galea (plurals ''laciniae'' and ''galeae''). At the outer margin, the typical galea is a cupped or scoop-like structure, located over the outer edge of the labium. In non-chewing insects, such as adult Lepidoptera, the maxillae may be drastically adapted to other functions. Unlike the mandibles, but like the labium, the maxillae bear lateral palps on their stipites. These palps serve as organs of touch and taste in feeding and in the inspection of potential foods and/or prey. In chewing insects, adductor and abductor muscles extend from inside the cranium to within the bases of the stipites and cardines much as happens with the mandibles in feeding, and also in using the maxillae as tools. To some extent the maxillae are more mobile than the mandibles, and the galeae, laciniae, and palps also can move up and down somewhat, in the
sagittal plane The sagittal plane (; also known as the longitudinal plane) is an anatomical plane that divides the body into right and left sections. It is perpendicular to the transverse and coronal planes. The plane may be in the center of the body and divid ...
, both in feeding and in working, for example in nest building by mud-dauber wasps. Maxillae in most insects function partly like mandibles in feeding, but they are more mobile and less heavily
sclerotised Sclerotin is a component of the cuticle of various Arthropoda, most familiarly insects. It is formed by cross-linking members of particular classes of protein molecules, a biochemical process called sclerotization, a form of tanning in which qu ...
than mandibles, so they are more important in manipulating soft, liquid, or particulate food rather than cutting or crushing food such as material that requires the mandibles to cut or crush. Like the mandibles, maxillae are innervated by the
subesophageal ganglia The suboesophageal ganglion (acronym: SOG; synonym: ''subesophageal ganglion'') of arthropods and in particular insects is part of the arthropod central nervous system (CNS). As indicated by its name, it is located ''below the'' ''oesophagus'', insi ...
.


Labium

The labium typically is a roughly quadrilateral structure, formed by paired, fused secondary maxillae. It is the major component of the floor of the mouth. Typically, together with the maxillae, the labium assists manipulation of food during
mastication Chewing or mastication is the process by which food is crushed and ground by teeth. It is the first step of digestion, and it increases the surface area of foods to allow a more efficient break down by enzymes. During the mastication process, ...
. The role of the labium in some insects however, is adapted to special functions; perhaps the most dramatic example is in the jaws of the
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ...
s of the Odonata, the dragonflies and
damselflies Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies, which constitute the other odonatan suborder, Anisoptera, but are smaller and have slimmer bodies. Most species fold the wings alo ...
. In these insects, the labium folds neatly beneath the head and thorax, but the insect can flick it out to snatch prey and bear it back to the head, where the chewing mouthparts can demolish it and swallow the particles. The labium is attached at the rear end of the structure called cibarium, and its broad basal portion is divided into regions called the submentum, which is the proximal part, the mentum in the middle, and the prementum, which is the distal section, and furthest anterior. The prementum bears a structure called the ligula; this consists of an inner pair of lobes called glossae and a lateral pair called paraglossae. These structures are homologous to the lacinia and galea of maxillae. The labial palps borne on the sides of labium are the counterparts of maxillary palps. Like the maxillary palps, the labial palps aid sensory function in eating. In many species the musculature of the labium is much more complex than that of the other jaws, because in most, the ligula, palps and prementum all can be moved independently. The labium is innervated by the sub-esophageal ganglia. In the honey bee, the labium is elongated to form a tube and tongue, and these insects are classified as having both chewing and lapping mouthparts. The wild silk moth ('' Bombyx mandarina'') is an example of an insect that has small labial palpi and no maxillary palpi.


Hypopharynx

The hypopharynx is a somewhat globular structure, located medially to the mandibles and the maxillae. In many species it is membranous and associated with salivary glands. It assists in swallowing the food. The hypopharynx divides the oral cavity into two parts: the cibarium or dorsal food pouch and ventral salivarium into which the salivary duct opens.


Siphoning insects

This section deals only with insects that feed by sucking fluids, as a rule without piercing their food first, and without sponging or licking. Typical examples are adult moths and butterflies. As is usually the case with insects, there are variations: some moths, such as species of ''
Serrodes ''Serrodes'' is a genus of moths in the family Erebidae first described by Achille Guenée in 1852. Description Costa of the forewings slightly arched before apex. Cilia of forewings and hindwings strongly crenulate. Tibia extremely hairy in b ...
'' and '' Achaea'' do pierce fruit to the extent that they are regarded as serious orchard pests. Some moths do not feed after emerging from the pupa, and have greatly reduced, vestigial mouthparts or none at all. All but a few adult Lepidoptera lack mandibles (the
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
known as the mandibulate moths have fully developed mandibles as adults), but also have the remaining mouthparts in the form of an elongated sucking tube, the proboscis.


Proboscis

The proboscis, as seen in adult Lepidoptera, is one of the defining characteristics of the morphology of the order; it is a long tube formed by the paired galeae of the maxillae. Unlike sucking organs in other orders of insects, the Lepidopteran proboscis can coil up so completely that it can fit under the head when not in use. During feeding, however, it extends to reach the nectar of flowers or other fluids. In certain specialist pollinators, the proboscis may be several times the body length of the moth.


Piercing and sucking insects

A number of insect orders (or more precisely families within them) have mouthparts that pierce food items to enable sucking of internal fluids. Some are herbivorous, like aphids and
leafhopper A leafhopper is the common name for any species from the family Cicadellidae. These minute insects, colloquially known as hoppers, are plant feeders that suck plant sap from grass, shrubs, or trees. Their hind legs are modified for jumping, and ...
s, while others are carnivorous, like assassin bugs and mosquitoes (females only).


Proboscis

The defining feature of the order Hemiptera is the possession of mouthparts where the mandibles and maxillae are modified into a proboscis, sheathed within a modified labium, which is capable of piercing tissues and sucking out the liquids. For example, true bugs, such as shield bugs, feed on the fluids of plants. Predatory bugs such as assassin bugs have the same mouthparts, but they are used to pierce the cuticles of captured prey.


Stylet

In female mosquitoes, all mouthparts are elongated. The labium encloses all other mouthparts like a sheath. The labrum forms the main feeding tube, through which blood is sucked. Paired mandibles and maxillae are present, together forming the stylet, which is used to pierce an animal's skin. During piercing, the labium remains outside the food item's skin, folding away from the stylet.
Saliva Saliva (commonly referred to as spit) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which DNA can ...
containing
anticoagulant Anticoagulants, commonly known as blood thinners, are chemical substances that prevent or reduce coagulation of blood, prolonging the clotting time. Some of them occur naturally in blood-eating animals such as leeches and mosquitoes, where t ...
s, is injected into the food item and blood sucked out, each through different tubes.


Sponging insects


Labellum

The
housefly The housefly (''Musca domestica'') is a fly of the suborder Cyclorrhapha. It is believed to have evolved in the Cenozoic Era, possibly in the Middle East, and has spread all over the world as a commensal of humans. It is the most common fl ...
is a typical sponging insect. The labellum's surface is covered by minute food channels, formed by the interlocking elongate hypopharynx and epipharynx, forming a proboscis used to channel liquid food to the oesophagus. The food channel draws liquid and liquified food to the oesophagus by capillary action. The housefly is able to eat solid food by secreting saliva and dabbing it over the food item. As the saliva dissolves the food, the solution is then drawn up into the mouth as a liquid.


References

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External links


Form & Function: the Insect HeadLabelled photos
Insect anatomy