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劳者秩序
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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。
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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
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]
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提出的可能辨识机制
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。
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]
马蚁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]
膜翅目例子
趋同演化例子：
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）。[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]
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马蚁 Pachycondyla inversa 会吃掉工蚁的卵（又称婚工蚁），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。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, 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
The tree wasp 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.[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 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, 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 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
无统治综合症
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
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. 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 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
In the 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]
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参考
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