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Mealworm
Temporal range:Holocene,0.003–0Ma
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Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Infraorder: Cucujiformia
Family: Tenebrionidae
Genus: Tenebrio
Species:
T. molitor
Binomial name
Tenebrio molitor
Mealworms (larvae ofTenebrio molitor) illustrated byDes Helmore

Mealwormsare the larval form of theyellow mealworm beetle,Tenebrio molitor,a species ofdarkling beetle.

The yellow mealworm beetle prefers a warmer climate and higher humidity.[1]Male mealworms release a sex pheromone to attract females to mate.[2]

Tenebrio molitorhas been used in biomedical research. Mealworms can also be a dietary source for animals and humans.[3]They are also considered pests, especially to food storage.

Description

Like allholometabolicinsects,T. molitorgoes through four life stages:egg,larva,pupa,andadult.Larvae typically measure about 2.5 centimetres (0.98 in) or more, whereas adults are generally 1.25 to 1.8 centimetres (0.49 to 0.71 in) in length.[1]

T. molitoris dark brown or black as an adult,[4]with larvae up to 1.25 inches (3.2 cm) long and adults up to 0.75 inches (1.9 cm) long.[5]

The yellow mealworm beetle can be differentiated from other beetles due to the linear grooves that are evenly divided and run along the abdomen. In addition, the beetle has only four tarsal segments on its hind legs. Most ground beetles, which are similar in size toTenebrio molitor,have five tarsal segments.[4]

Commonly mistaken with the black mealworm beetle (T. obscurus), key variations in regard to size and shape distinguish these two. The abdomen of the adult black mealworm beetle is more rounded and ends in a pointed tip, as opposed to the more rectangular and blunt-ended abdomen of the yellow mealworm beetle.[4]Additionally, the larvae ofT. molitorare lighter colored than those ofT. obscurus.[5]

Distribution

Mealworms most-likely originated in theMediterraneanregion, but are now present in many areas of the world as a result of human trade and colonization. The oldest archaeological records of mealworms can be traced toBronze AgeTurkey.Records from the British Isles and northern Europe are from a later date, and mealworms are conspicuously absent from archaeological finds from ancientEgypt.[6]

Life cycle

The eggs hatch 4 to 19 days after the female oviposits.[7]

During the larval stage, the mealworms feed on vegetation and dead insects andmoltbetween each larval stage, orinstar(9 to 20 instars). After the final molt, theypupate.The new pupa is whitish and turns brown over time. After 3 to 30 days, depending on environmental conditions such as temperature, it emerges as an adult beetle.[7]

As larvae

In studies,T. molitorlarvae show an incubation period of seven to eight days and a period of three to four days for the first instar. After the first instar, there is significant variation for the number of days in each instar period, though variation may be due to malnutrition orpathogens.Before emergence, most larvae typically go through 15 to 17 instars, with very few larvae going through the 19 to 20 instars.[8]The body length of the larvae gradually increases with each successive instar, reaching maximum length at the 17th instar. However, the body length decreases beyond the 17th instar. Pupation occurs after the 14th instar, with most larvae showing totalpupationbetween the 15th and 17th instars. Additionally, larvae are white in the first instar and gradually turn brown after the second instar.[8]

Effect of parental age and temperature

Offspring produced by older beetles have shorter larval stages than those produced by younger beetles. Larvae from the older beetles also show a rapid weight increase at an earlier age than those from young parents. At 25°C, the larval stage was shortened, the number of larval molts decreased, and the durations of adult life decreased when parental age increased compared to the beetles at 30°C.[9]Another study found that at 20°, 25°, and 30°C, parental age does not have any effect on the duration of the egg stage or the weights of the eggs.

However, the amount of hatched eggs decreased when parental age increased. When eggs were laid during the first two months after emergence, approximately 90% of the eggs hatched, but when they were laid after four months, only about 50% hatched.[9]It was also found that larvae from young parents grow at a slower rate compared to larvae produced by the same parents nine weeks earlier. At 30°C, there were no other effects of parental age on the larvae. However, at 20° and 25°C, the larvae from young parents required significantly more time to complete development and had more molts compared to the larvae from the same parents after they had aged one more or longer. Additionally, the duration of adult life decreased when parental age increased.[9]

Reproduction

Tenebrio molitorlarvae eating an apple slice

A sexpheromonereleased by male mealworms has been identified.[10]Inbreedingreduces the attractiveness of sexual pheromone signaling by male mealworms.[2]Females are more attracted to the odors produced byoutbredmales than the odors produced by inbred males. The reduction of male signaling capability may be due to increased expression of homozygous deleterious recessive alleles caused by inbreeding.[11]

The mealworm beetle breeds prolifically. Males insert sperm packets with theiraedeagus.Within a few days, the female burrows into soft ground and lays eggs. Over her adult lifespan of about 6-12 months, a female will, on average, lay about 500 eggs.[7]

Mating

Researchers examined whether female beetles preferred noninfected or infected males as mates. Virgin females were found to spend more time near and copulated more often with noninfected males in the experiment. Mating behavior was also found to be influenced by the mass of the male beetle. A larger male was preferred and mated with first compared to a smaller male mate. Overall, female beetles showed mating preferences for noninfected males who were larger in size. One explanation provided by researchers for this findings is that noninfected males who are larger may allow females to gain genetic or material benefits.[12]

Immunocompetence

Evidence suggests that in many animal species, secondary sexual traits reflect male immunocompetence, the ability of an individual’s immune system to resist and control pathogens or parasites. A study found that a single parasite-like immunological challenge, created via a nylon monofilament implant in the beetle, significantly reduced the sexual attractiveness and locomotor activity of males, it did not negatively affect their survival. When the inserts were removed, the majority of the males showed greater encapsulation responses of the implant, though some of the males seemed to have already chosen a terminal reproductive investment strategy.[13]And thus, the majority of males invest in their immune system after the first challenge.

A second immune challenge increased their attractiveness, but was found to significantly reduce locomotor activity of the males and increase their mortality. This represents a trade-off between pheromone production and energy required for activities such as immune system recovery and locomotor activity.[13]

When there was a third challenge (implantation) in the same males, there was a lower encapsulation rate of the nylon implants in more attractive males than the less attractive, showing how the males made no attempts to boost their immune system.[13]The results suggest that the males who become sexually attractive after the second immune challenge have a trade-off where they sacrifice locomotor activity and do not energetically invest in immune system recovery. This shows how the female mealworm beetles consistently preferred males who invested significantly less in immune system recovery and that males are not able to allocate resources simultaneously to both improving their health, or in this case, recovery of their immune system, and to increasing their sexual attractiveness.[13]

Nutritional condition

Pheromonesare chemical signals that function as mate attractors and relay important information to prospective mates. For a reliable signal, it must be costly to produce, which means it is likely to have condition dependent expression. A study found that female preference of the pheromones was dependent on the nutritional condition of the males, shown by how they spent significantly more time with males who received constant food than males who received no food.[14]It was also found thatphenoloxidaseactivity was dependent on the nutritional condition of the males, with phenoloxidase activity being two to six times higher in males with constant food than in males who received no food. However, nutritional conditions had no effect on the encapsulation rate of the males.[14]

When receiving constant food, male initial body mass had no correlations with phenoloxidase activity or encapsulation rate. This shows that pheromone mediated attractiveness and the immunocompetence in terms of phenoloxidase activity of males were condition-dependent, as both decreased with nutritional stress. This suggests that there is a trade-off between allocation of resources and energy into the production of pheromones and immunocompetence and that the production of pheromones are condition dependent sexual traits.[14]

Immune defense

The mealworm beetle is currently considered a pest when infesting and degrading the quality of stored grains or grain products. However, they are now being promoted as a beneficial insect, as their high nutrient content makes them a viable food source for pet food, protein-rich animal feed, or even human nutrition, and they are capable of degrading plastic waste and polystyrene.[15]These benefits make mealworms attractive for mass rearing, a technique that promotes disease transmission within the colonies.T. molitorcan be the host of many different pathogens and parasites, including entomopathogenicmicrobes,protozoa,andtapeworms,which can decrease the mealworm beetle’s survival or reproductive success.[15]

Behavioral immunity

When the mealworms feed on infected rodent feces, they may consume the eggs of the tapeworm parasiteHymenolepis diminuta.Infected male beetles pay a higher reproductive cost than the female beetles.T. molitordisplays behavioral immunity when exposed toH. diminuta,shown by how the infected males develop an avoidance behavior for feces that harbor the tapeworm, which decreases their probability of coming into contact with the tapeworm in the future.[15]The female mealworm beetles develop a qualitative resistance through mate choice, as they are able to evaluate male immunocompetence through pheromone signaling, allowing them to choose the more immunologically fit males as mates. This also reduces the probability of the females being infected by their mates and may cause them to pass on an enhanced level of immunocompetence to their offspring.[15]

Another way mealworms may display behavioral immunity is how they may tolerate infections by limiting negative effects on their reproductive success. For example, mealworm beetles tolerate a high number of cysticercoids ofH. diminutaat the expense of their own fitness and longevity. But the males produce improved spermatophores that contain superior nuptial gifts that will be passed to their mating females, increasing female fecundity and causing a greater number of eggs to be fertilized.[15]

Cuticular color

Cuticular color, a heritable component, of the mealworm beetle varies from tan to black. In the mealworm beetle, evidence suggests that population level variation in cuticular color is linked to pathogen resistance in that darker individuals are more resistant to pathogens. A study found that two immune parameters related to resistance, haemocyte density and pre-immune challenge activity of phenoloxidase, were significantly higher in selection lines of black beetles compared to tan lines. Higher haemocyte density is likely indicative of a heightened immune response.[16]

There were no effects of gender on the immune traits. Cuticular color is dependent on melanin production, which requires phenoloxidase, an enzyme that is present in its inactive form inside haemocytes. This shows why darker insects have a heightened immune response and are more resistant to pathogens that invade the hemocoel via the cuticle. However, there was no significant difference in haemolymph antibacterial activity between black and tan lines, explained by how antimicrobial peptides are produced by haemocytes but are not involved in cuticular darkening.[16]

InT. molitor,the degree of cuticular melanization is a strong indicator of resistance to the entomopathogenic fungusMetarhizium anisopliae,which could be explained by the thicker and less porous cuticle displayed by darker insects compared to lighter ones. However, there seems to be underlying trade-offs that prevent the fixation of the darker phenotype, shown by how the plasticity of melanization phenotypes in response to population density may contribute to the absence of predominance of darker individuals amongT. molitorpopulations.[15]

Food restriction

Because immune defenses against parasites and pathogens require metabolic resources, food restriction can impair immune function ofT. molitor.For adultT. molitorbeetles, phenoloxidase activity can be reduced by half during short-term food deprivation but returns rapidly to original levels when the beetles are given access to food again.[15]Additionally,T. molitorlarvae can eat five times more food per day than usual following an immune challenge to compensate for the caloric loss from the immune response. These immune challenged larvae also show significant weight loss when fed with either protein or carbohydrate rich diets but show stable weights when given both protein and carbohydrate-rich diets.[15]

HealthyT. molitorlarvae usually prefer diets with a lower protein to carbohydrate ratio but shift toward food with higher protein contents after an immune challenge with bacteria. This causes enhanced hemocyte circulation and antibacterial activity in the hemolymph, which likely maximizes resistance against bacterial infection. However, phenoloxidase activity is not affected by this shift in diet choice.[15]

A study found that the effects of nutritional imbalance on body composition were buffered by the subsequent selection of complementary foods. This demonstrates that the mealworm beetles are capable of compensating for nutritional imbalances and that the way nutritional balance is restored depends on the nutrient that is initially deficient in their food.[17]For example, if the beetles were previously fed a protein-rich, carbohydrate-deficient diet, they would prefer carbohydrates to protein, whereas beetles fed a carbohydrate-rich, protein-deficient diet, they would strongly prefer a protein-rich diet. They found that self-selectingT. molitorbeetles recovered from carbohydrate or protein deficiency within six days by selecting the complementary diet.[17]

Gut microbiota

The gut microbiota ofT. molitorconsists of many different bacterial species present at low abundance. A study found a predominance of genusSpiroplasmaspecies in the phylum Tenericutes in the gut samples ofT. molitor,but there was variation found in the community composition between individuals. Although someSpiroplasmaspecies are known insect pathogens, theT. molitorlarvae did not experience any harmful effects from the presence of theSpiroplasmain the gut, indicating that they are not pathogens to the host.[18]By comparing this to the bacterial communities found in other insects, it was found that theSpiroplasmaspecies found were specific toT. molitor.Additionally, the gut bacteria community structure was not significantly affected by the presence of antibiotics or by the exposure of the beetle larvae to a more highly diverse soil bacteria community. There is a negative relationship between bacterial diversity and ampicillin concentration, meaning ampicillin treatment caused a reduction in the bacterial community size, which was determined with pyrosequencing of the 16S rRNA gene, and no negative relationship when kanamycin was added.[18]

Polystyrene foamdecreasesT. molitorfecundity, but the beetle can fully develop using the plastic as its primary source of food, which makes it an interesting alternative to recycle polystyrene. However, when the mealworm’s microbiota is disrupted by an antibiotic treatment, it loses its ability to digest polystyrene, suggesting that its associated gut microbes are essential in the digestion process. Specifically, the bacteriumExiguobacteriumspecies, which was isolated from the midgut of mealworms, was demonstrated to degrade polystyrene in vitro.[15]

Relationship with humans

As feed and pet food

Main articles:Insects as feedandInsect based pet food

Mealworms are typically used as apet foodfor captivereptiles,fish,birds,and some smallmammals.They are also provided to wild birds inbird feeders,particularly during the nesting season. Mealworms are useful for their high protein content. They are also used asfishing bait.[19]

They are commercially available in bulk and are typically available in containers withbranoroatmealfor food. Commercial growers incorporate ajuvenile hormoneinto the feeding process to keep the mealworm in the larval stage and achieve an abnormal length of 2 cm or greater.[20]

As food

Main articles:Insects as foodandEntomophagy in humans
Mealworms in a bowl

Mealworms areedible for humans,and processed into several insect food items available in food retail such as insect burgers.[21]

Due to their high protein and fat content, as well as consuming large amounts of fiber, they represent a good food source for humans. They are also high in oleic acid, which can decreaselow-density lipoprotein(LDL) and increasehigh-density lipoprotein(HDL) levels in the blood.[8]

Mealworms have historically been consumed in many Asian countries, particularly in Southeast Asia. There, they are commonly found in food markets and sold as street food alongside other edible insects. Baked or fried mealworms have been marketed as a healthy snack food in recent history, though the consumption of mealworms goes back centuries.[3]

In May 2017, mealworms were approved as food in Switzerland.[22]In June 2021, dried mealworms were authorized asnovel foodin theEuropean Union,[23]after theEuropean Food Safety Authorityassessed the larvae as safe for human consumption.[24][25]

Mealworm larvae contain significant nutrient content.[19]For every 100 grams of raw mealworm larvae, 206 kilocalories and anywhere from 14 to 25 grams of protein are contained.[26]Mealworm larvae contain levels of potassium, copper, sodium, selenium, iron and zinc that rival those of beef. Mealworms contain essential linoleic acids as well. They also have greater vitamin content by weight compared to beef,B12not included.[26][27]

Mealworms may be easily reared on fresh oats, wheat bran or grain, with sliced potato, carrots, or apple as a moisture source. The small amount of space required to raise mealworms has made them relevant for scalable industrialized mass production.[28]

Possible health risks

Some evidence suggests thatT. molitormay pose a health risk, as humans and animals can consume the eggs and larvae of the beetle with grain-based food. Although they are usually either digested or are excreted with feces, sometimes, they are able to survive and live in thealimentary tract.The first cases ofT. molitorlarvae in human organs date back to the 19th century, where their presence was observed in thegastrointestinal tract,including thestomachandintestines.[29]There were other cases, such as a reported ulcer infestation ofT. molitorin anAIDSpatient and a concerned urinary canthariasis in a ten year old boy in Iran in 2019, which was the last reported human case of canthariasis caused byT. molitor.However, there are very few reported cases of live larvae in animals, and there are no reports of gastrointestinal canthariasis in farm animals.[29]

In waste disposal

In 2015, it was discovered that 100 mealworms can degradepolystyreneinto usable organic matter at a rate of about 34–39 milligrams per day. Additionally, no difference was found between mealworms fed only Styrofoam and the mealworms fed conventional foods, during the one-month duration of the experiment.[30]Microorganisms inside the mealworm's gut are responsible for degrading the polystyrene, with mealworms given theantibioticgentamicinshowing no signs of degradation.[31]Isolated colonies of the mealworm's gut microbes, however, have proven less efficient at degradation than the bacteria within the gut.[31]

See also

  • Zond 5,a 1968 space mission on which mealworms were among the first terrestrial organisms to travel to and circle the Moon[32]
  • Plastivore

Gallery

  • In a bedding of bran
    In a bedding of bran
  • Mealworm detail
    Mealworm detail
  • A mealworm pupa with molted larval skin
    A mealworm pupa with molted larval skin
  • New adult
    New adult
  • Mature adult
    Mature adult

References

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  2. ^abPölkki M, Krams I, Kangassalo K, Rantala MJ (2012)."Inbreeding affects sexual signalling in males but not females ofTenebrio molitor".Biology Letters.8(3): 423–5.doi:10.1098/rsbl.2011.1135.PMC3367757.PMID22237501.
  3. ^abMaster, The Party (2022-06-28)."EDIBLE MEALWORMS: EVERYTHING You Need to Know!".Party Bugs – Seasoned party snacks made from edible insects (crickets and mealworms).Retrieved2022-09-30.
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  5. ^abBennett, Stuart M. (2003)."Tenebrio molitor(Yellow Mealworm Beetle) ".Stored Product Insects.Retrieved2024-03-20.
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  13. ^abcdKrams, I; Daukšte, J; Kivleniece, I; Krama, T; Rantala, Mj; Ramey, G; Šauša, L (2011-01-11)."Female Choice Reveals Terminal Investment in Male Mealworm Beetles, Tenebrio molitor, after a Repeated Activation of the Immune System".Journal of Insect Science.11(56): 56.doi:10.1673/031.011.5601.ISSN1536-2442.PMC3281432.PMID21864151.
  14. ^abcRantala, Markus J.; Kortet, Raine; Kotiaho, Janne S.; Vainikka, Anssi; Suhonen, Jukka (2003)."Condition Dependence of Pheromones and Immune Function in the Grain Beetle Tenebrio molitor".Functional Ecology.17(4): 534–540.Bibcode:2003FuEco..17..534R.doi:10.1046/j.1365-2435.2003.00764.x.ISSN0269-8463.JSTOR3598991.
  15. ^abcdefghijVigneron, Aurélien; Jehan, Charly; Rigaud, Thierry; Moret, Yannick (2019)."Immune Defenses of a Beneficial Pest: The Mealworm Beetle, Tenebrio molitor".Frontiers in Physiology.10:138.doi:10.3389/fphys.2019.00138.ISSN1664-042X.PMC6422893.PMID30914960.
  16. ^abArmitage, S. a. O.; Siva-Jothy, M. T. (2005-06-01)."Immune function responds to selection for cuticular colour in Tenebrio molitor".Heredity.94(6): 650–656.doi:10.1038/sj.hdy.6800675.ISSN1365-2540.PMID15815710.
  17. ^abRho, Myung Suk; Pum LEE, Kwang (2015-04-02)."Nutrient-specific food selection buffers the effect of nutritional imbalance in the mealworm beetle, Tenebrio molitor (Coleoptera: Tenebrionidae)".European Journal of Entomology.112(2): 251–258.doi:10.14411/eje.2015.030.
  18. ^abJung, Jaejoon; Heo, Aram; Park, Yong Woo; Kim, Ye Ji; Koh, Helium; Park, Wooden (2014-05-27)."Gut Microbiota of Tenebrio molitor and Their Response to Environmental Change"(PDF).Journal of Microbiology and Biotechnology.24(7): 888–897.doi:10.4014/jmb.1405.05016.PMID24861345.
  19. ^abRavzanaadii, Nergui; Kim, Seong-Hyun; Choi, Won-Ho; Hong, Seong-Jin; Kim, Nam-Jung (2012-09-30)."Nutritional Value of Mealworm, Tenebrio molitor as Food Source".International Journal of Industrial Entomology.25(1): 93–98.doi:10.7852/ijie.2012.25.1.093.ISSN1598-3579.
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  23. ^"Commission Implementing Regulation (EU) 2021/882 of 1 June 2021 authorising the placing on the market of driedTenebrio molitorlarva as a novel food under Regulation (EU) 2015/2283 of the European Parliament and of the Council, and amending Commission Implementing Regulation (EU) 2017/2470 ".EU Commission. 2 June 2021.
  24. ^Turck, Dominique; Castenmiller, Jacqueline; De Henauw, Stefaan; Hirsch-Ernst, Karen Ildico; Kearney, John; MacIuk, Alexandre; Mangelsdorf, Inge; McArdle, Harry J.; Naska, Androniki; Pelaez, Carmen; Pentieva, Kristina; Siani, Alfonso; Thies, Frank; Tsabouri, Sophia; Vinceti, Marco; Cubadda, Francesco; Frenzel, Thomas; Heinonen, Marina; Marchelli, Rosangela; Neuhäuser-Berthold, Monika; Poulsen, Morten; Prieto Maradona, Miguel; Schlatter, Josef Rudolf; Van Loveren, Henk; Ververis, Ermolaos; Knutsen, Helle Katrine; Knutsen, H. K. (13 January 2021)."Safety of dried yellow mealworm (Tenebrio molitorlarva) as a novel food pursuant to Regulation (EU) 2015/2283] ".EFSA Journal.19(1): 6343.doi:10.2903/j.efsa.2021.6343.PMC7805300.PMID33488808.
  25. ^Boffey, Daniel (2021-01-13)."Yellow mealworm safe for humans to eat, says EU food safety agency".The Guardian.ISSN0261-3077.Retrieved2021-01-15.
  26. ^ab"6. nutritional value of insects for human consumption"(PDF).Edible insects: future prospects for food and feed security(PDF).FAO FORESTRY PAPER. Vol. 171. FAO. 2013. pp. 67–.
  27. ^Schmidt, Anatol; Call, Lisa; Macheiner, Lukas; Mayer, Helmut K. (2018). "Determination of vitamin B12 in four edible insect species by immunoaffinity and ultra-high performance liquid chromatography".Food Chemistry.281:124–129.doi:10.1016/j.foodchem.2018.12.039.PMID30658738.S2CID58651702.
  28. ^Rumbos, Christos I.; Athanassiou, Christos G. (March 2021)."Insects as Food and Feed: If You Can't Beat Them, Eat Them!'—To the Magnificent Seven and Beyond".Journal of Insect Science.21(2): 9.doi:10.1093/jisesa/ieab019.PMC8023366.PMID33822126.
  29. ^abGałęcki, Remigiusz; Michalski, Mirosław Mariusz; Wierzchosławski, Karol; Bakuła, Tadeusz (2020-11-11)."Gastric canthariasis caused by invasion of mealworm beetle larvae in weaned pigs in large-scale farming".BMC Veterinary Research.16(1): 439.doi:10.1186/s12917-020-02657-0.ISSN1746-6148.PMC7659124.PMID33176765.
  30. ^Jordan, Rob (29 September 2015)."Plastic-eating worms may offer solution to mounting waste, Stanford researchers discover".Stanford News Service.Archived fromthe originalon 20 February 2021.Retrieved24 March2016.
  31. ^abLockwood, Deirdre (September 30, 2015)."Mealworms Munch Polystyrene Foam".Chemical and Engineering News.Retrieved2019-02-04.
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