| Manduca sexta | |
|---|---|
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| Caterpillar | |
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| Adult moth | |
Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Lepidoptera |
| Family: | Sphingidae |
| Genus: | Manduca |
| Species: | M. sexta |
| Binomial name | |
| Manduca sexta | |
| Synonyms | |
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Manduca sexta is amoth of the familySphingidae present through much of theAmericas. The species wasfirst described byCarl Linnaeus in his 1763Centuria Insectorum.
Commonly known as theCarolina sphinx moth and thetobacco hawk moth (as adults) and thetobacco hornworm and theGoliath worm (as larvae), it is closely related to and often confused with the very similartomato hornworm (Manduca quinquemaculata); thelarvae of both feed on the foliage of various plants of the familySolanaceae. The larvae of these species can be distinguished by their lateral markings: Tomato hornworms have eight V-shaped white markings with no borders; tobacco hornworms have seven white diagonal lines with a black border. Additionally, tobacco hornworms have red horns, while tomato hornworms have dark blue or black horns.[2] Amnemonic to remember the markings is tobacco hornworms have straight white lines like cigarettes, while tomato hornworms have V-shaped markings (as in "vine-ripened" tomatoes).M. sexta has mechanisms for selectively sequestering and secreting theneurotoxinnicotine present in tobacco.[citation needed]
M. sexta is a commonmodel organism, especially inneurobiology, due to its easily accessiblenervous system and short life cycle. Due to its immense sizeM. sexta is big enough for medical imaging modalities (likeCT,MRI, orPET) and used as a model in imaging and gut inflammation.[3] It is used in a variety of biomedical and biological scientific experiments. It can be easily raised on a wheat-germ-based diet. The larva is large, and thus it is relatively easy to dissect it and isolate its organs.
M. sexta has a short life cycle, lasting about 30 to 50 days. In most areas,M. sexta has about two generations per year, but can have three or four generations per year in Florida.[4]
M. sextaeggs are spherical, approximately 1.5 millimeters in diameter, and translucent green.[2] They typically hatch two to four days after they are laid. Eggs are normally found on the underside of foliage, but can also be found on the upper surface.
M. sexta larvae are bright green in color and grow up to 100 millimeters in length. The posteriorabdominal segment is tipped with a dorsocaudal horn that earns them the name "hornworm". The finalinstar consists of a cylindrical body covered with fine hairlikesetae. The head is equipped with a pair ofocelli andchewing mouthparts. Each of the threethoracic segments bears a pair of true legs, and there is a pair ofprolegs on the third, fourth, fifth, sixth, and last abdominal segments in all larval instars. Theprothoracic segment bears one pair ofspiracles, and additional pairs occur on each of the eight abdominal segments.[5]
Thehemolymph (blood) of this species contains insecticyanin, a blue-coloredbiliprotein. When the larva feeds on its normal diet of plant foliage, it ingestspigmentaciouscarotenoids, which are primarily yellow in hue. The resulting combination is green. Under laboratory conditions—when fed a wheat-germ-based diet—larvae are turquoise in color due to the lack of carotenoids in their diet.[citation needed]
Thecaterpillar stage of the tobacco hornworm is quite similar in appearance to that of the closely relatedtomato hornworm. The larvae of these two species can however be readily distinguished by their lateral markings. Specifically, theM. sexta caterpillar has seven white diagonal lines with a black border at the first seven abdominal segments, and the horn is red or green with a red tip. TheM. quinquemaculatacaterpillar has V-shaped white markings with no borders at all eight of its abdominal segments, and the horn is dark blue or black in color.[6]
During the larval stage,M. sexta caterpillars feed on plants of the familySolanaceae, principally tobacco, tomatoes and members of the genusDatura.M. sexta has five larval instars, which are separated byecdysis (molting), but may add larval instars when nutrient conditions are poor. Near the end of this stage, the caterpillar seeks a location forpupation, burrows underground, and pupates. This searching behavior is known as "wandering". The imminence of pupation—suggested behaviorally by the wandering—can be anatomically confirmed by spotting theheart (aorta), which is a long, pulsating vessel running along the length of the caterpillar's dorsal side. The heart becomes visible through the skin just as the caterpillar is reaching the end of the final instar.
A commonbiological control for hornworms is theparasiticbraconid waspCotesia congregata, which lays its eggs in the bodies of the hornworms. The wasp larvae feed internally and emerge from the body to spin theircocoons. Parasitized hornworms are often seen covered with multiple white, cottony wasp cocoons, which are often mistaken for large eggs. Awasp species,Polistes erythrocephalus, feeds on hornworm larvae.[7]
Before the larva pupates, it goes through a stage called the pre-pupa, where it shrinks considerably and prepares to pupate. Often people mistake this stage for a dead or dying caterpillar.
The pupal stage lasts approximately 14–18 days under laboratory conditions (17 hours light, 7 hours dark, 27 °C). When reared on a short-dayphotoperiod (12 hours light, 12 hours dark), pupae enter a state ofdiapause that can last several months. During the pupal stage, structures of the adult moth form within the pupal case, which is shed duringeclosion (adult emergence).
AdultM. sexta have narrow wings with a wing span of approximately 100 mm.M. sexta moths arenectarivorous and feed on flowers, demonstrating a remarkable ability to hover.
Adults aresexually dimorphic. Males are identifiable by their broaderantennae and the presence ofclaspers at the end of the abdomen. Female moths are typically ready to mate one week aftereclosion, and do so only once. Males may mate many times. Mating generally occurs on a vertical surface at night, and can last several hours, with the male and female facing in opposite positions, their posterior ends touching. After mating, females deposit their fertilized eggs on foliage, usually on the underside of leaves.
LikeDrosophila melanogaster,M. sexta is commonly used as amodel organism for experiments. They are frequently studied in the laboratory due to their large size and relative ease of rearing. They may be reared on host plants, such as tobacco and tobacco relatives, tomato plants, or wheat-germ-based artificial diet. Their rearing is straightforward, provided they receive a long daylight cycle (e.g., 14 hours) during development to preventdiapause.
Eggs are rinsed for one to five minutes in dilute household bleach for disinfection.
Eggs are placed on diet cubes or host plants. The eggs hatch and develop at different speeds depending on temperature. The larvae are moved to a fresh diet or leaves as their food spoils or is consumed. When they start to "wander", they are about to pupate, so are placed in a pupation chamber. Pupation chambers are holes drilled into a wood board. TheManduca larvae are sealed in the chamber using a stopper and allowed to pupate. After pupation, the pupae are placed in a breeding or colony chamber toeclose. Providing a cup of sugar water and a tobacco (or related) plant will allow mated females tooviposit fertile eggs, which can then be reared.
When fed an artificial diet,Manduca larvae do not consume thexanthophyll -which is a yellow pigment- needed to produce their green coloration; instead they appear blue. On some diets, they have very little pigment and pigment precursors, so are a very pale blue-white. As vitamin A and othercarotenoids are necessary for the visual pigments (rhodopsin), an artificial-diet-reared hornworm may have poor vision due to lack of carotenoids in the diet.[8]
Captive-bred hornworms fed on an artificial diet are often given to insectivorous exotic animals, such as certain reptiles, fish and small mammals. They are preferred over wild-collected hornworms, which maybioaccumulate poisonous substances found in dietary plants. Hornworms, though originally bred for laboratories, are also farmed for this purpose.[9][10][11] They are often sold already packed into pods that include everything the larvae need, including food. Care is relatively easy, and animals seem to relish their bright color and flavor.[12]
M. sexta larvae grow up to 100 millimeters in length, reaching up to 20 grams. Due to their large size, they are used as alternative animal models for medical imaging modalities likecomputed tomography,magnetic resonance imaging, orpositron emission tomography.[13] Researchers aroundAnton Windfelder have established the larvae ofM. sexta as an alternativeanimal model for chronicinflammatory bowel diseases or as an animal model for testing newcontrast agents forradiology.[14]
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Tobacco hornworms are facultative specialists; the larvae can grow and develop on any host plants. However, the larvae prefer solanaceous plants, such as tobacco and tomato plants. On these types of plants, larvae grow and develop faster. The lateral and medialsensilla styloconia (sensory receptors) on their mouthparts help them to identify solanaceous plants by recognizingindioside D, a steroidal glycoside found in those particular plants (del Campo et al., 2001).[15][16]Salicin is a distastefulphagodeterrent, found only inSalix spp. whilecaffeine is a phagodeterrent that is actually toxic.[16] Schoonhoven 1969 found thatM. sextahabituation to salicin is mediated bydesensitization of the deterrence associated peripheral neurons and Glendinning et al. 1999 the same for caffeine.[16] However Glendinning et al. 2001 find only a small peripheral desensitization for salicin, concluding that Schoonhoven erred, and that habituation in this case is centrally mediated.[16] Tobacco hornworms are considered pests because they feed on the upper leaves of tobacco plants and leave green or black droppings on the plants. As adults, they do not damage plants since they feed on nectar.[17]
Tobacco hornworm larvae prefer humid environments. When dehydrated, tobacco hornworm larvae will move towards a source of water or to an area with a high relative level of humidity. They use their antennae to locate water to drink .[18]
Nicotine is poisonous to most animals that use muscles to move because nicotine targets theacetylcholine receptor at theneuromuscular junction. However, the tobacco hornworm is capable of metabolizing nicotine from the tobacco plant and using nicotine as a defense against predators. It possesses a gene called cytochrome P450 6B46 (CYP6B46) that converts nicotine into a metabolite. About 0.65% of nicotine metabolites are transported from the gut to thehemolymph, where they are reconverted to nicotine and released into the air from the tobacco hornworm's spiracles. The emitted nicotine is used as a way to deter spiders, a practice known as “toxic halitosis.” In one study, tobacco hornworms that fed from nicotine-deficient plants or expressed low levels of CYP6B46 were more susceptible to wolf spider predation.[19]
Tobacco hornworm caterpillars emit short clicking sounds from their mandibles when they are being attacked. This sound production is believed to be a type of acousticaposematism, or warning sounds that let predators know that trying to eat them will be troublesome; tobacco hornworms have been observed to thrash and bite predators after producing those clicking sounds. These clicks can be heard at a close distance with a frequency range of 5 to 50 kHz. The intensity of clicks increases with the number of attacks (Bura et al., 2012).[20]
