维基百科

劳者秩序

劳者纠察、工者整饬、工者辖制（英文：Worker policing，直译：工者治安、劳动者管辖）是社会性膜翅目动物（蚂蚁、蜜蜂、黄蜂）常见的行为：雌性工蚁会吃掉其它工蚁产的卵，确保蚁后的后代在群体占据主导。在某些蜜蜂、蚂蚁、黄蜂物种，工蜂或蜂王也可能直接攻击有生育能力的工蜂。

单倍二倍体性别决定系统的理论基础
这存在于蜜蜂和其它膜翅目昆虫中，包括大黄蜂、蚂蚁、黄蜂等等。

这种强制的纠察或促进真社会性社会的利他演化。[1]

其可能通过表面标记的碳氢化合物或角质层碳氢化合物识别。[2][3][4] 极少数情况，工蜂会携带模仿蚁后的碳氢化合物逃避监管，这种情况又称“无统治综合症”、“无秩序综合症”（英文：anarchic syndrome）。[5]

也并非所有纠察都需要女王；一些蚂蚁就建立了雌蚁统治等级，就让排名高的个体繁殖。[6]


演化基础

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参见：染色体倍性

大多社会性昆虫透过染色体倍性性别决定系统决定性别。单倍体雄性由未受精卵发育而来，而二倍体雌性则由受精卵发育而来。许多蚂蚁、蜜蜂、黄蜂物种，工蜂工蚁虽然无法交配，但仍保留有功能的卵巢。[7]因此，它们能产生雄性后代。

平均而言，女王与她的儿子能分享一半基因，但与女工的儿子们却只分享四分之一的基因。因此，抚养自己的儿子而不是女工的儿子才符合女王的最大利益。[7]

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双方最大化其生殖成就导致王和工在贡献不同雄性后代比例。女王喜欢生产自己的儿子，女工则喜欢生产自己的儿子，并且不会因为与侄子有良好的亲缘关系而阻止姐妹产蛋。[7]

Worker policing occurs when worker bees in the colony are genetically more closely related to the queen than the reproductive female. In many instances, the eggs of the female worker are eaten, or she is repeatedly attacked by other workers. This mechanism of egg removal ensures that the queen's sons predominate.[8]

Experiments confirming the role of kin selection in worker policing demonstrate the effects of multiple matings, which can lead to lower average relatedness between workers. Queens of the social wasp Dolichovespula saxonica mate singly or multiply. Researchers who study these wasps have observed a strong positive correlation between worker relatedness and male production. After controlling for the absolute number of eggs laid, these scientists conclude that the queen's multiple matings favor mutual worker policing.[9]

Although early theories of worker policing focused on the kin selection component, evidence from honey bees suggests that altruism is sometimes enforced. Fewer workers reproduce as policing effectiveness rises, and policing effectiveness decreases with increasing relatedness except in colonies with no queen. This suggests that worker policing is a social sanction imposed on selfish individuals.[10]

提出的可能辨识机制

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Formica fusca蚂蚁可以辨识巢友

可能包括女王的碳氢化合物、女工的碳氢化合物或巢友识别。

女王的碳氢化合物

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对弓背蚁属 Camponotus floridanus的研究发现，其女王的卵表有特殊的碳氢化合物。在有女王产卵的群体，工蚁不产卵；而没有女王产卵的群体，工蜂则繁殖。因此，碳氢化合物还可提醒工蚁停止繁殖。 目前还不清楚限制繁殖如何发生，或者减少工蜂繁殖所需的最低讯号阈值是多少。[2]

女工的碳氢化合物

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马蚁Platythyrea punctata（英语：Platythyrea punctata）的群体中，角质层化合物的存在引发了蚁群内部攻击，特别是新旧繁殖工蚁之间。具体来说，老的生殖工蚁会在新雌性的触角涂抹一种标记碳氢化合物，将它们识别为竞争对手和攻击目标。[3]

同窝识别

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黑山蚁似乎还包括巢友识别。更多攻击非同巢工蚁，并带走其蛋。[4]

膜翅目例子

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趋同演化例子：

蜜蜂

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西方蜜蜂吃掉工蚁的卵（卵食（英语：Oophagy））

最早发现的是西方蜜蜂（Apis mellifera）。普遍工蜂管制让工蜂繁殖率极低（0.12%）。[11] 蜜蜂透过吃来维持繁殖率（卵食（英语：Oophagy））。[8]许多蜂巢工蜂卵巢活性减弱，产卵受抑制。 [8]

据报道，小蜜蜂（Apis florea）也有卵食（英语：Oophagy）行为。透过微卫星分析，没有成熟的雄蜂具有非蜂王等位基因。因此，即使工蚁的卵巢已活化并且能够产卵，工蚁的监管也确保了工蚁的功能性不孕。[12]


据报导，东方蜜蜂雌性工蜂激活卵巢的比例高于意大利蜜蜂或小蜜蜂。当蜂后移除后，高达40%工蜂几天内激活自己卵巢。然而，工蜂继续吃工蚁的卵，显示该物种的工者辖制并不涉及蜂后。[13]

对一妻多夫制（英语：Polyandry in animals）蜜蜂属来说也是类似。[13]

劳者秩序也发生在原始的社会性熊蜂，例如欧洲熊蜂。[14]

蚂蚁

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Harpegnathos saltator（英语：Harpegnathos saltator）杀害外来女王

马蚁 Pachycondyla inversa 会吃掉工蚁的卵（又称婚工蚁（英语：gamergate (ant)）），并攻击产卵的蚁。[15]

在 Gnamptogenys menadensis 的工蚁有时会交配产卵，而其它工蚁产用作食物的营养卵（英语：trophic egg）。由于其工蚁既可产雄性卵，也可产雌性卵，因此青睐劳者秩序。其劳者秩序可能有固定女工六肢（据报半数死亡），也可能拖出蚁群。[16]

Aphaenogaster smythiesi japonica 也显示出劳者秩序的证据。当研究人员将重新组合有蚁后蚁群和无蚁后蚁群，工蚁会攻击卵巢激活的工蚁。[17]

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In the queenless ant Streblognathus peetersi, policing helps to select gamergates. The workers immobilize gamergates with lowered fertility, enabling high-ranking workers to take over as new gamergates.[18]

In the ponerine ant Harpegnathos saltator（英语：Harpegnathos saltator）, worker policing prevents the number of reproductive workers from becoming excessive. The species has gamergates, which lay eggs but rarely carry out any other tasks, so having too many of them would reduce the colony's efficiency. Gamergate numbers are rather stable, as infertile workers target newly ovipositing workers by jumping on and holding them: this efficiently inhibits them from laying eggs, and within a few weeks they revert to being subordinate and infertile workers. However, once new ovipositing workers attain a gamergate's level of ovarian activity, the infertile (policing) workers ignore them. The signal used by the policing workers consists of cuticle hydrocarbons transmitted only by direct contact, rather than being a 信息素 detected by 嗅觉. Similarly, the inhibition of ovipositing workers is achieved by direct physical aggression by policing workers, not by any pheromone produced by queens or gamergates.[19]

Wasps

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Dolichovespula sylvestris（英语：Dolichovespula sylvestris） workers can display aggression towards other fertile workers

The tree wasp Dolichovespula sylvestris（英语：Dolichovespula sylvestris） also displays worker policing. Both egg-eating and aggression are reported in these wasp communities, and the queen also engages in the policing process.[20] Similar behavior has been observed in the closely related species Dolichovespula media（英语：Dolichovespula media）.[21]

In colonies of the paper wasp, 中华马蜂, workers can lay up to a quarter of the male eggs in the colony. The number of eggs that survived to hatching, however, was minimal compared to the number produced by the queen. Analysis of the microsatellite markers showed that both queens and workers contributed to policing of worker and queen-laid eggs even in Monogyny（英语：Monogyny） and monandrous colonies.[22]

The common wasp, 普通黄胡蜂, engages in worker policing, and it is known that a significant number of workers have active ovaries. However, studies by researchers have suggested that relatedness may not be the key factor in the development of worker policing. Rather, it is hypothesized that worker policing has been selected due to conflict suppression in the colony.[23]

In a test of worker policing, researchers examined the removal of worker-laid eggs in Vespula rufa（英语：Vespula rufa）, a wasp that has low paternity. Under the testing conditions, worker policing was not as efficient, and some of the drones appeared to be from worker-laid eggs.[24]

The 黄边胡蜂, Vespa crabro, was previously thought to be under reproductive pheromone control by the queen, thus explaining why the other females did not reproduce, even though they were capable of doing so. Experiments by Foster showed that the workers were instead regulating sterility in each other, thus instead exhibiting worker policing. While each worker is capable of reproducing, the colony as a whole is more efficient and organized if the workers allow only the queen to lay larvae.[25]

Synoeca cyanea（英语：Synoeca cyanea） also engage in worker policing during times when queen repopulation is not needed. They will prevent females from laying eggs through aggressive behavior and egg-eating.[26]

例外

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无统治综合症

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在极少数情况下，雌性工蜂的卵可以逃脱监管，这个过程被称为无政府症候群。

Very rarely, female worker bees lay eggs that escape worker policing, in a process known as the anarchic syndrome. Female reproductive workers in these colonies activate their ovaries even in the presence of a queen. The worker eggs are allowed as they 拟态 queen hydrocarbons. Thus, female workers can maximize their reproductive fitness at the expense of the colony. Anarchic syndrome is an example of selection working in opposite directions at individual and group levels.[5]

自私劳者秩序

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Temnothorax unifasciatus（英语：Temnothorax unifasciatus）, a 家蚁亚科 ant, has been shown not to have collective worker policing. However, when a queen is removed, a rank order for reproduction appears, where top ranking reproductives display aggression towards lower-order female workers. Thus, reductions in egg laying are created by Dominance hierarchy（英语：Dominance hierarchy）. Individuals do not act in benefit of the colony, instead opting to increase their own fitness by laying their eggs and reducing the contributions of opponents.[6]

This type of policy is common in Dolichovespula norwegica（英语：Dolichovespula norwegica） wasps where despite being more related to other workers' sons, worker-laid eggs were consumed by other workers.[27]

没基因冲突的劳者秩序

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In the Thelytoky（英语：Thelytoky） ant Platythyrea punctata, colonies are 克隆; therefore, workers are all equally related genetically, and worker policing is not expected as there is no genetic conflict. However, one would expect severe effects on the condition of the colony if reproductive workers were left unchecked. For example, more time spent on individual reproduction would be predicted to detract from care of the queen's offspring. In studies of these ants, workers did actually display worker policing, as evidenced by increased aggression towards reproductive workers. Worker policing here is favored for group efficiency.[28]

参考

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  1. ^ Ratnieks, Francis L.W.; Heikki Helanterä. The evolution of extreme altruism and inequality in insect societies. Phil. Trans. R. Soc. B. October 2009, 364 (1553): 3169–3179. PMC 2781879  . PMID 19805425. doi:10.1098/rstb.2009.0129. 
  2. ^ 2.0 2.1 Endler, Annett; Jürgen Liebig; Bert Hölldobler. Queen fertility, egg marking and colony size in the ant Camponotus floridanus. Behavioral Ecology and Sociobiology. February 2006, 59 (4): 490–499. S2CID 22084023. doi:10.1007/s00265-005-0073-0. 
  3. ^ 3.0 3.1 Hartmann, Anne; Patrizia D'Ettorre; Graeme R. Jones; Jürgen Heinze. Fertility signaling—the proximate mechanism of worker policing in a clonal ant. Naturwissenschaften. June 2005, 92 (6): 282–286. Bibcode:2005NW.....92..282H. PMID 15770464. S2CID 11123443. doi:10.1007/s00114-005-0625-1. 
  4. ^ 4.0 4.1 Helanterä, Heikki; Liselotte Sundström. Worker policing and nest mate recognition in the ant Formica fusca. Behavioral Ecology and Sociobiology. June 2007, 61 (8): 1143–1149. S2CID 9920557. doi:10.1007/s00265-006-0327-5. 
  5. ^ 5.0 5.1 Oldroyd, Benjamin P.; Katherine E Osborne. The evolution of worker sterility in honeybees: the genetic basis of failure of worker policing. Proc. R. Soc. Lond. B. July 1999, 266 (1426): 1335–1339. PMC 1690071  . doi:10.1098/rspb.1999.0784. 
  6. ^ 6.0 6.1 Stroeymeyt, Nathalie; Elisabeth Brunner; Jürgen Heinze. "Selfish worker policing" controls reproduction in a Temnothorax ant. Behavioral Ecology and Sociobiology. July 2007, 61 (9): 1449–1457. S2CID 3191624. doi:10.1007/s00265-007-0377-3. 
  7. ^ 7.0 7.1 7.2 Ratnieks, Francis L.W.; P. Kirk Visscher. Worker policing in the honeybee. Nature. December 1989, 342 (6251): 796–797. Bibcode:1989Natur.342..796R. S2CID 4366903. doi:10.1038/342796a0. 
  8. ^ 8.0 8.1 8.2 Wenseleers, Tom; Adam G. Hart; Francis L. W. Ratnieks. When Resistance Is Useless: Policing and the Evolution of Reproductive Acquiescence in Insect Societies (PDF). The American Naturalist. December 2004, 164 (6): E154–E167 [2024-03-02]. JSTOR 10.1086/425223. PMID 29641925. S2CID 4809642. doi:10.1086/425223. （原始内容存档 (PDF)于2024-03-03）. 
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  11. ^ Visscher, P. Kirk. A quantitative study of worker reproduction in honey bee colonies. Behavioral Ecology and Sociobiology. 1989, 25 (4): 247–254. JSTOR 4600337. S2CID 37903505. doi:10.1007/bf00300050. 
  12. ^ Halling, Luke A; Benjamin P. Oldroyd; Wandee Wattanachaiyingcharoen; Andrew B. Barron; Piyamas Nanork; Siriwat Wongsiri. Worker policing in the bee Apis florea. Behavioral Ecology and Sociobiology. 2001, 49 (6): 509–513. S2CID 9930323. doi:10.1007/s002650100325. 
  13. ^ 13.0 13.1 Oldroyd, Benjamin P.; Luke A. Halling; Gregory Good; Wandee Wattanachaiyingcharoen; Andrew B. Barron; Piyamas Nanork; Siriwat Wongsiri; Francis L. Ratnieks. Worker policing and worker reproduction in Apis cerana. Behavioral Ecology and Sociobiology. 2001, 50 (4): 371–377. S2CID 15558324. doi:10.1007/s002650100376. 
  14. ^ Zanette, L. R.; Miller, S. D.; Faria, C. M.; Almond, E. J.; Huggins, T. J.; Jordan, W. C.; Bourke, A. F. Reproductive conflict in bumblebees and the evolution of worker policing. Evolution. December 2012, 66 (12): 3765–3777 [2024-03-02]. PMID 23206135. S2CID 36787898. doi:10.1111/j.1558-5646.2012.01709.x. （原始内容存档于2023-06-26）. 
  15. ^ D'Ettorre, Patrizia; Jürgen Heinze; Francis L W Ratnieks. Worker policing by egg eating in the ponerine ant Pachycondyla inversa. Proc Biol Sci. July 2004, 271 (1546): 1427–1434. PMC 1691738  . PMID 15306343. doi:10.1098/rspb.2004.2742. 
  16. ^ Gobin, Bruno; J. Billen; C. Peeters. Policing behaviour towards virgin egg layers in a polygynous ponerine ant. Anim. Behav. November 1999, 58 (5): 1117–1122. PMID 10564615. S2CID 16428974. doi:10.1006/anbe.1999.1245. 
  17. ^ Iwanishi, Satoru; Eisuke Hasegawab; Kyohsuke Ohkawaraa. Worker oviposition and policing behaviour in the myrmicine ant Aphaenogaster smythiesi japonica Forel. Animal Behaviour. September 2003, 66 (3): 513–519. S2CID 28820297. doi:10.1006/anbe.2003.2222. 
  18. ^ Gadau, Jürgen; Fewell, Jennifer; Wilson, Edward O. Organization of Insect Societies: From Genome to Sociocomplexity. Harvard University Press. 2009: 227–228. ISBN 978-0-674-03125-8. 
  19. ^ Liebig, Jürgen; Peeters, Christian; Höllldobler, Bert. Worker policing limits the number of reproductives in a ponerine ant. Proc. R. Soc. Lond. B. 1999, 266 (1431): 1865–1870. PMC 1690207  . doi:10.1098/rspb.1999.0858. 
  20. ^ Wenseleers, Tom; A. Tofilski; F. L. W. Ratnieks. Queen and worker policing in the tree wasp Dolichovespula sylvestris. Behavioral Ecology and Sociobiology. 2005, 58: 80–86. S2CID 16813247. doi:10.1007/s00265-004-0892-4. 
  21. ^ Foster, Kevin R. Colony kin structure and male production in Dolichovespula wasps. Molecular Ecology. 2001, 10 (4): 1003–1010. PMID 11348506. S2CID 12009153. doi:10.1046/j.1365-294X.2001.01228.x. 
  22. ^ Saigo, T.; K. Tsuchida. Queen and worker policing in monogynous and monandrous colonies of a primitively eusocial wasp. Proc Biol Sci. December 2004, 271 (Suppl 6): S509–S512. PMC 1810092  . PMID 15801618. doi:10.1098/rsbl.2004.0238. 
  23. ^ Foster, Kevin R.; Francis L. W. Ratnieks. Convergent evolution of worker policing by egg eating in the honeybee and common wasp. Proc Biol Sci. January 2001, 268 (1463): 169–74. PMC 1088587  . PMID 11209887. doi:10.1098/rspb.2000.1346. 
  24. ^ Wenseleers, Tom; N. S. Badcock; K. Erven; A. Tofilski; F. S. Nascimento; A. G. Hart; T. A. Burke; M. E. Archer; F. L W. Ratnieks. A test of worker policing theory in an advanced eusocial wasp, Vespula rufa. Evolution. June 2005, 59 (6): 1306–14. PMID 16050107. S2CID 15644821. doi:10.1554/04-532. 
  25. ^ Foster, K.R., Gulliver, J., Ratnieks, F.L.W. Worker policing in the European hornet Vespa crabro. Insectes Sociaux. 2002, 49 (1): 41–44. S2CID 45182961. doi:10.1007/s00040-002-8277-z. 
  26. ^ Noda, Silvia Cristina Mari. Morphological and Physiological Caste Differences in Synoeca cyanea (Hymenoptero, Vespidae, Epiponini) According to the Ontogenetic Development of the Colonies. Sociobiology. 2003. 
  27. ^ Bonckaert, W.; Tofilski, A.; Nascimento, F.S.; Billen, J.; Ratnieks, F.L.W.; Wenseleers, T. Co-occurrence of three types of egg policing in the Norwegian wasp Dolichovespsula wasp. Behavioral Ecology and Sociobiology. 2001, 65 (4): 633–640. S2CID 2186614. doi:10.1007/s00265-010-1064-3. 
  28. ^ Hartmann, Anne; J. Wantia; J.A. Torres; J. Heinze. Worker policing without genetic conflicts in a clonal ant. PNAS. October 2003, 100 (22): 12836–12840. Bibcode:2003PNAS..10012836H. PMC 240705  . PMID 14557542. doi:10.1073/pnas.2132993100  . 

外部链接

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检索自“https://zh.wikipedia.org/w/index.php?title=勞者秩序&oldid=83274628”