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Article Evaluation

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I chose to evaluate the article called "Fixed action pattern". This article has received a quality rating of “start class”, meaning it does not provide a strong over view of the topic. In fact, there are multiple ways this article could be improved.

The lead paragraph “Fixed action pattern” (FAP) is very brief. The definition of the topic is adequate in explaining the meaning of the term FAP but the rest of the overview is lacking detail. In particular, the 6 characteristics identified by Lorenz of what make a behaviour a FAP are listed in the lead paragraph without any explanation of what these terms mean. These characteristics should have been described in more detail to offer the readers a more complete understanding of the topic.

The rest of the structure is clear, with headings organized by theme. However, I believe the order of the sections should be rearranged with the “Examples” section listed last, after the topic has been thoroughly discussed. The sections in the article have even coverage, but it would be beneficial to allocate more prominence to the description and explanation of the topic in the introduction. Also, the “Terminology” section is not necessary. The terms discussed in this section could be included in the lead paragraph because they would be useful in making the overview more complete. The “Examples” section could be improved by explaining the examples listed under “Others” in more depth and in relation to the topic, instead of just listing them. The examples should also be described in closer relation to the FAP, the stimuli and 6 characteristics of FAP to ensure there is no confusion amongst the readers on how the examples demonstrate the topic. The “Significance” section could be improved by discussing the evolutionary significance and the development of FAP behaviours in terms of evolutionary advantages more clearly and in more depth. The “Exploitation” section could also be included in the “Significance” section, opposed to being its own vague section. This is because it only mentions briefly how mimicry is used to induce FAP, but it could be looked at from a point of how mimicry of sign stimuli and releasers offer evolutionary advantages and it could be discussed in terms of this significance. The article could also benefit from the addition of more sections to help the readers understand the topic. For example, there could be a history section to discuss the origin of studying FAP, Wallace Craig’s Consummatory Behaviour as a starting point, other early scientists who studied FAP and the significance of the emergence of this idea in Classical Ethology.

This article is written in a neutral perspective, which is encouraged on Wikipedia, however, there is no mention of exceptions or different perspectives such as a comparative psychology perspective; which would look at the impact of learning on these behaviours. They could include an “Exceptions” section in order to give a more holistic view of this area of research.

The sources used in this article are reliable, with most of them coming from University presses and textbooks. However there are very few references used, making it hard to check the facts. Many facts are not referenced including many of the examples, statements, terminology definitions and facts. For example, there are no citations in “Terminology” section and most of the examples given in the “Examples” section are also not cited. There is also reference to an unnamed source of information in the statement “FAPs have been said to…” which is frowned upon by Wikipedia.

On the talk page there are multiple discussions about what examples qualify as FAP behaviours. This may relate to the unclear explanations of the examples used. There have also been 2 sections removed from the article due to poor examples used and unreliable sources.

           Our discussion on FAP in class differed from this article in a few ways. We began our discussion with an understanding of the basis of this field as we learned about classical ethology, Wallace Craig’s components of behaviour (not mentioned in this article) and the transition of Craig’s Consummatory behaviour idea to Lorenz’s FAP idea. These are important steps to the emergence of FAP research, which were not well represented in the article. We then discussed what FAPs are in detail, putting emphasis on Lorenz’s 6 characteristics of FAP in order to fully understand this type of behvaiour, which the article failed to do. We then went on to discuss examples that were clearly explained in relation to the FAP behaviour, the stimuli that elicit them and how it fits the 6 characteristics of FAP. Many of the same examples were discussed in class as in the article, but the article did not do a sufficient job in highlighting how the examples fit the topic.

Multiple other details could be improved in this article such as:

·     The grammar in some sentences needs correction

·     The first Lorenz should be hyperlinked as opposed to the second

·     Supernormal stimuli should be explained where it was mentioned

·     A photograph of the Greylag Goose demonstrating egg rolling should be used as opposed to a picture of just the goose

Overall this article did use correct information but should be enhanced in many areas.

Week 4: Add to an Article

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Article: Weaving (Horse).

Added cited definition in the lead section.

Added cited information to the "Causes" section.

Week 5: Choose possible topics

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Article: Weaving (Horse). Start class.

To improve:

-Add to lead: Explain its relation to abnormal animal behaviour and stereotypic animal behaviour.

-Add to causes section: There are more causes than mentioned. Explain exceptions.

-Add missing citations.

-Add pictures.

TA: This sounds like a good topic, just make sure there's enough published literature to support your additions and you may want to consider adding a new section (unless you feel confident you can make a significant contribution by adding to the existing sections).

Article: Trophallaxis. Start class.

Week 6: Draft Article

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Article: Trophallaxis

This article is a start class article of mid importance.

The limited information already present in the article is reliable and cited, with multiple examples provided of animals that engage in this behaviour. Therefore, I plan to keep this information but reorganize it in a logical format, as well as add new information. I will create new sections (stated below) to organize the existing information, as well as additional information and examples to explain the topic in more detail.

I will add to the definition of Trophallaxis in the lead paragraph: along with nutrients, trophallaxis can involve the transfer of molecules and organisms such as pheromones and symbionts, as well as information to serve as a form of communication (Suarez & Throne, 2014). The lead will not include any information past the first paragraph already existing, as the rest of the information will be moved to other appropriate sections.

I will move the section Etymology to the top of the article, just below the lead. This section will include the origin of the word, as well as the sentence already present regarding the introduction of the word by William Morton Wheeler. Sections to add:

Evolutionary Significance: This will include the evolutionary advantages of food sharing, and some of the fitness benefits associated with the behaviour. I will include some examples, such as significance of food sharing in bats from the article by Carter & Wilkinson (listed below). I may also incorporate various purposes of the behaviour, as they relate back to adaptive significance such as: nutrition, immunity, foraging communication, kin survival, etc.

Mechanisms (or some variant heading): This will include mechanisms and physiology of food transfer. I will add examples, such as the mechanism described in honey bees by Gil & De Marco, as well as the reciprocal sharing in wasps, described by Suryanarayanan & Jeanne (sources listed below).

Trophallaxis in Insects: This will include the examples provided on the page already, since the majority of them are insects. I will include the purpose and significance of the behaviour in each example. I will also add examples I have found, such as the transfer of immunity through trophallaxis in termites (Hamilton, Lejeune, & Rosengaus, 2011) and associative learning through Trophallaxis in honey bees (Gil & De Marco, 2005).

Trophallaxis in Vertebrates: This will include the 2 examples of vertebrates (wolves and bats) already in the article, however I will find support for these examples and explain the behaviours in more detail for these species. I will also find more examples of vertebrates that engage in trophallaxis, such as birds. This section is significant to demonstrate the behaviour is found in a diverse set of species, not only insects.

Overall, I plan to create 5 sections in the article, fully explaining the mechanisms and significance of trophallaxis as it is found in different species.

Bibliography:

The following sources will be used to add to the article, plus additional ones I find as I continue to research the topic and develop the entire article.

  • Suarez, M. E., & Throne, B. L. (2000). Rate, Amount, and Distribution Pattern of Alimentary Fluid Transfer via Trophallaxis in Three Species of Termites (Isoptera: Rhinotermitidae, Termopsidae). Annals of the Entomological Society of America, 93(1),145-155. https://doi.org/10.1603 http://www.bioone.org/doi/abs/10.1603/0013-8746(2000)093%5B0145:RAADPO%5D2.0.CO%3B2/0013-8746(2000)093[0145:RAADPO]2.0.CO;2   
  • Carter, G., & Wilkinson, G. (2013). Does food sharing in vampire bats demonstrate reciprocity?Communicative & Integrative Biology. 6(6), https://doi-org.qe2a-proxy.mun.ca/10.4161/cib.25783
  • Gil, M., & De Marco, R. J. (2005). Olfactory learning by means of trophallaxis in Apis mellifera. Journal of Experimental Biology, 208, 671-680. doi: 10.1242/jeb.01474
  • Suryanarayanan, S., & Jeanne, R. L. (2008). Antennal Drumming, Tophallaxis and Colony Development in the Social Wasp Polistes fascatus (Hymenoptera: Vespidae). Ethology, 114, 1201–1209. doi:10.1111/j.1439-0310.2008.01561.x
  • Hamilton, C., Lejeune, B. T., & Rosengaus, R. B. (2011). Trophallaxis and prophylaxis: social immunity in the carpenter ant Camponotus pennsylvanicus. Biology Letters, 7(1), 89–92. http://doi.org/10.1098/rsbl.2010.0466

Week 10: First Draft

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Trophallaxis /ˌtroʊfəlˈæksɪs/ is the transfer of food or other fluids among members of a community through mouth-to-mouth (stomodeal) or anus-to-mouth (proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pheromones, organisms such as

Trophallaxis in Asian-Australian weaver ant O. smaragdina, Thailand

symbionts, and information to serve as a form of communication.[1] Trophallaxis is most highly developed in social insects such as ants, termites, wasps and bees.

Etymology

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Tropho- (prefix or suffix) is derived from the Greek trophé, meaning 'nourishment'. The Greek 'allaxis' means 'exchange'.[2] The word trophallaxis was introduced by the entomologist William Morton Wheeler in 1918.[3]

Evolutionary significance

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Trophallaxis was used in the past to support theories on the origin of sociality in insects.[4][5] The Swiss psychologist and entomologist Auguste Forel also believed that food sharing was key to ant society and he used an illustration of it as the frontispiece for his book The Social World of the Ants Compared with that of Man.[6] Besides sociality, trophallaxis has evolved within many species as a method of nourishment for adults and/ or juveniles,[7] kin survival, [7] transfer of symbionts,[8] transfer of immunity,[9] colony recognition[10] and foraging communication.[11] Trophallaxis has even evolved as a parasitic strategy in some species to obtain food from their host.[12] Trophallaxis can also result in the spreading of chemicals, such as pheromones, throughout a colony; which is significant in social colony functioning.[13] Species have evolved anatomy to allow them to participate in trophallaxis, such as the proventriculus in the crops of Formica Fusca ants.[13] This structure acts as a valve to enhance food storage capacity.[13] Likewise, the honey bee Apis mellifera is able to protrude their proboscis and sip nectar from the open mandibles of the donor bee.[11] Certain mechanisms have also evolved to initiate food sharing, such as the sensory exploitation strategy that has evolved in the common cuckoo brood parasites.[14] These birds have evolved brightly coloured gapes that stimulate the host to transfer food.[14]

Trophallaxis in invertebrates

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Tophallaxis is a form of social feeding in many insects that contributes to the formation of social bonds.[5] Trophallaxis serves as a means of

Trophallaxis in Camponotus sp.

communication, at least in bees, like M. genalis.[15] Trophallaxis in M. genalis is part of a social exchange system, where dominant bees are usually the recipients of food.[15] It increases longeivity of bees that have less access to food and decreases aggression between nest mates.[15] In the red fire ant,[16] colony members store food in their crops and regularly exchange this food with other colony members and larvae to form a sort of "communal stomach" for the colony. This is also true for certain species of Lasioglossum, such as the sweat bee Lasioglossum hemichalceum. L. hemichalceum will often exchange food with other members regardless of whether they are nestmates or not.[17] This is because cooperation among non-relatives offers more benefit than cost to the group.[17]

Many wasps, like Protopolybia exigua and Belonogaster petiolata, exhibit foraging behavior where adults perform trophallaxis with adults and between adults and larvae.[18][19] P. exigua carry nectar, wood pulp and macerated prey in its crop from the field to the nest for transfer; for larvae survival they carry amounts of prey proportional to the amount of larvae in the nest.[18] Voluntary trophallaxis in Xylocopa pubescens bees has led to the nest guarding behavior that the species is known for.[20] This bee species allows one adult to forage and bring nectar back for the rest of the nest population as a way to continually defend the nest while obtaining nutrients for all members of the colony.

In termites,[21] proctodeal trophallaxis is crucial for replacing the gut endosymbionts that are lost after every molt. Gut symbionts are also transferred by anal trophallaxis in wood-eating termites.[22] This should not be confused with coprophagia. Transfer of gut symbionts in these species is essential to digest wood as their food source. Carpenter ants transfer immunity through trophallaxis by the direct transfer of antimicrobial substances, increasing disease resistance and social immunity of the colony.[9] [20]

In some species of ants, it may play a role in spreading the colony odour that identifies members.[23]

Honey bee foragers use trophallaxis in associative learning to form long-term olfactory memories, in order to teach nest mates foraging behaviour and where to search for food. [11]

In addition, Vespula austriaca wasps also engage in trophallaxis as a form of parasitism with its host workers to obtain nutrients.[24] V. austriaca is an obligate parasite species that invades the nests of host species and obtains food by constraining the host with their legs and forcing trophallaxis.[24]

Trophallaxis in vertebrates

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Food sharing between parent and chick Pygoscelis antarctica penguins

Vertebrates such as some bird species, gray wolves, and vampire bats feed their young through reguritation of food as a form of trophallaxis. Food sharing in vertebrates is a form of reciprocity demonstrated by many social vertebrates.[25]

Wild wolves transport food in their stomach to pups and/or breeding females and share it by regurgitation, as a form of trophallaxis.[26] The recipient wolves often lick or sniff the donor wolf's muzzle to activate regurgitation and receive nutrients.[26] Vampire bats share blood with kin by regurgitation as a means of increasing their fitness through kin selection.[25]

Birds regurgitate food and directly transfer it into the mouths of their offspring as a part of parental care, such as the "crop milk" that is transferred by mother ring doves into the mouths of their young.[27] The cuckoo brood parasite is another bird species that engages in trophallaxis. The cuckoo bird uses mimicry, such as mimicking the egg shell colours and patterns of the host's eggs, to place their young in the nest of host species where they will be fed and reared at no expense to the cuckoo mother.[14] The cuckoo young can often mimic the begging call of an entire nest of the host species' young and have evolved intensely coloured gaits; which both act as supernormal stimuli to induce the host bird to deliver food via trophallaxis.[14]

  1. ^ E., Suárez, Mark; L., Thorne, Barbara (2000-01-01). "Rate, Amount, and Distribution Pattern of Alimentary Fluid Transfer via Trophallaxis in Three Species of Termites (Isoptera: Rhinotermitidae, Termopsidae)". Annals of the Entomological Society of America. 93 (1). doi:10.1603/0013-8746(2000)093[0145:RAADPO]2.0.CO;2. ISSN 0013-8746.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  2. ^ "trophallaxis | Definition of trophallaxis in US English by Oxford Dictionaries". Oxford Dictionaries | English. Retrieved 2018-03-01.
  3. ^ Wheeler, W. M. (1918). "A study of some ant larvae with a consideration of the origin and meaning of social habits among insects". Proc. Am. Philos. Soc. 57 (4): 293–343. JSTOR 983940.
  4. ^ Roubaud, E. (1916). "Recherches biologiques sur les guepes solitaires et sociales d'Afrique. La genese de la vie sociale et l'evolution de l'instinct maternel chez les vespides". Ann Sci Nat (in French). 1: 1–160.
  5. ^ a b Wcislo, W. T.; Gonzalez, V. H. (2006-05-01). "Social and ecological contexts of trophallaxis in facultatively social sweat bees, Megalopta genalis and M. ecuadoria (Hymenoptera, Halictidae)". Insectes Sociaux. 53 (2): 220–225. doi:10.1007/s00040-005-0861-6. ISSN 0020-1812.
  6. ^ Sleigh, Charlotte (2002). "Brave new worlds: Trophallaxis and the origin of society in the early twentieth century". Journal of History of the Behavioral Sciences. 38 (2): 133–156. doi:10.1002/jhbs.10033.
  7. ^ a b Keeping, Malcolm G. (1997-03-01). "Social behavior and brood decline in reproductive-phase colonies ofBelonogaster petiolata (Degeer) (Hymenoptera: Vespidae)". Journal of Insect Behavior. 10 (2): 265–278. doi:10.1007/BF02765559. ISSN 0892-7553.
  8. ^ Kitade, Osamu (2004). "Comparison of Symbiotic Flagellate Faunae between Termites and a Wood-Feeding Cockroach of the Genus Cryptocercus". Microbes and environments. 19 (3): 215–220. doi:10.1264/jsme2.19.215. ISSN 1342-6311.
  9. ^ a b Hamilton, Casey; Lejeune, Brian T.; Rosengaus, Rebeca B. (2011-02-23). "Trophallaxis and prophylaxis: social immunity in the carpenter ant Camponotus pennsylvanicus". Biology Letters. 7 (1): 89–92. doi:10.1098/rsbl.2010.0466. ISSN 1744-957X. PMC 3030872. PMID 20591850.{{cite journal}}: CS1 maint: PMC format (link)
  10. ^ Dahbi, Abdallah; Hefetz, Abraham; Cerdá, Xim; Lenoir, Alain (1999-07-01). "Trophallaxis Mediates Uniformity of Colony Odor in Cataglyphis iberica Ants (Hymenoptera, Formicidae)". Journal of Insect Behavior. 12 (4): 559–567. doi:10.1023/A:1020975009450. ISSN 0892-7553.
  11. ^ a b c Gil, Mariana; Marco, Rodrigo J. De (2005-02-15). "Olfactory learning by means of trophallaxis in Apis mellifera". Journal of Experimental Biology. 208 (4): 671–680. doi:10.1242/jeb.01474. ISSN 0022-0949. PMID 15695759.
  12. ^ Reed, H. C.; Akre, R. D. (1983-09-01). "Colony behavior of the obligate social parasiteVespula austriaca (Panzer) (Hymenoptera: Vespidae)". Insectes Sociaux. 30 (3): 259–273. doi:10.1007/BF02223984. ISSN 0020-1812.
  13. ^ a b c Chapman, R. F. (2013). The Insects Structure and Function. New York, USA: Cambridge University Press. p. 42. ISBN 978-0-521-11389-2.
  14. ^ a b c d "Bird brood parasitism". Current Biology. 23 (20): R909–R913. 2013-10-21. doi:10.1016/j.cub.2013.08.025. ISSN 0960-9822.
  15. ^ a b c Wcislo, W. T.; Gonzalez, V. H. (2006-05-01). "Social and ecological contexts of trophallaxis in facultatively social sweat bees, Megalopta genalis and M. ecuadoria (Hymenoptera, Halictidae)". Insectes Sociaux. 53 (2): 220–225. doi:10.1007/s00040-005-0861-6. ISSN 0020-1812.
  16. ^ "Allocation of liquid food to larvae via trophallaxis in colonies of the fire ant, Solenopsis invicta". Animal Behaviour. 50 (3): 801–813. 1995-01-01. doi:10.1016/0003-3472(95)80140-5. ISSN 0003-3472.
  17. ^ a b Kukuk, P. F.; Sage, G. K. (1994-12-01). "Reproductivity and relatedness in a communal halictine beeLasioglossum (Chilalictus) hemichalceum". Insectes Sociaux. 41 (4): 443–455. doi:10.1007/BF01240647. ISSN 0020-1812.
  18. ^ a b Rocha, AA, and Carlos Costa Bichara Filho. "Resources Taken to the Nest by Protopolybia exigua (Hymenoptera, Vespidae) in Different Phases of the Colony Cycle, in a Region of the Medio Sao Fransisco River, Bahia, Brazil."Sociobiology54.2 (2009): 439–56. Print.https://www.researchgate.net/publication/261833590_Resources_Taken_to_the_Nest_by_Protopolybia_exigua_Hymenoptera_Vespidae_in_Different_Phases_of_the_Colony_Cycle_in_a_Region_of_the_Medio_Sao_Francisco_River_Bahia_Brazil
  19. ^ Keeping, Malcolm G. "Social Behavior and Brood Decline in Reproductive-phase Colonies Of Belonogaster Petiolata(Degeer) (Hymenoptera: Vespidae)." Journal of Insect Behavior 10.2 (1997): 265–78. Web. 20 Sept. 2014.https://link.springer.com/article/10.1007/BF02765559
  20. ^ a b van der Blom, Jan; Velthuis, Hayo H. W. (1988-01-12). "Social Behaviour of the Carpenter Bee Xylocopa pubescens (Spinola)". Ethology. 79 (4): 281–294. doi:10.1111/j.1439-0310.1988.tb00717.x. ISSN 1439-0310.
  21. ^ Kitade, Osamu (2004). "Comparison of Symbiotic Flagellate Faunae between Termites and a Wood-Feeding Cockroach of the Genus Cryptocercus". Microbes and Environments. 19 (3): 215–220. doi:10.1264/jsme2.19.215.
  22. ^ Klass, Klaus-Dieter; Nalepa, Christine; Lo, Nathan (March 2008). "Wood-feeding cockroaches as models for termite evolution (Insecta: Dictyoptera): Cryptocercus vs. Parasphaeria boleiriana". Molecular Phylogenetics and Evolution. 46 (3): 809–817. doi:10.1016/j.ympev.2007.11.028. ISSN 1055-7903. PMID 18226554.
  23. ^ Dahbi, A.; Hefetz, A.; Cerda, X.; Lenoir, A. (1999). "Trophallaxis mediates uniformity of colony odor in Cataglyphis iberica ants (Hymenoptera, Formicidae)". Journal of Insect Behavior. 12 (4): 559–567. doi:10.1023/A:1020975009450.
  24. ^ a b Reed, H.C.; Akre, R.D. (1983). "Colony behavior of the obligate social Vespula austriaca (Panzer) (Hymenoptera: Vespidae)". Insectes Sociaux. 30 (3): 259–273. doi:10.1007/bf02223984.
  25. ^ a b Carter, Gerald; Wilkinson, Gerald (2013-11-01). "Does food sharing in vampire bats demonstrate reciprocity?". Communicative & Integrative Biology. 6 (6). doi:10.4161/cib.25783. ISSN 1942-0889. PMC 3913674. PMID 24505498.{{cite journal}}: CS1 maint: PMC format (link)
  26. ^ a b Mech, L David; Wolf, Paul C; Packard, Jane M (1999-11-01). "Regurgitative food transfer among wild wolves". Canadian Journal of Zoology. 77 (8): 1192–1195. doi:10.1139/z99-097. ISSN 0008-4301.
  27. ^ Lehrman, Daniel S. (1955-01-01). "The Physiological Basis of Parental Feeding Behavior in the Ring Dove (Streptopelia Risoria)". Behaviour. 7 (1): 241–285. doi:10.1163/156853955X00094. ISSN 1568-539X.
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