勞者秩序

勞者糾察、工者整飭、工者轄制（英文：Worker policing，直譯：工者治安、勞動者管轄）是社會性膜翅目動物（螞蟻、蜜蜂、黃蜂）常見的行為：雌性工蟻會吃掉其它工蟻產的卵，確保蟻后的後代在群體佔據主導。在某些蜜蜂、螞蟻、黃蜂物種，工蜂或蜂王也可能直接攻擊有生育能力的工蜂。

這種強制的糾察或促進真社會性社會的利他演化。

其可能通過表面標記的碳氫化合物或角質層碳氫化合物識別。
極少數情況，工蜂會攜帶模仿蟻后的碳氫化合物逃避監管，這種情況又稱「無統治綜合症」、「無秩序綜合症」（英文：anarchic syndrome）。

也並非所有糾察都需要女王；一些螞蟻就建立了雌蟻統治等級，就讓排名高的個體繁殖。

 演化基礎 

大多社會性昆蟲透過染色體倍性性別決定系統決定性別。單倍體雄性由未受精卵發育而來，而二倍體雌性則由受精卵發育而來。許多螞蟻、蜜蜂、黃蜂物種，工蜂工蟻雖然無法交配，但仍保留有功能的卵巢。因此，它們能產生雄性後代。

平均而言，女王與她的兒子能分享一半基因，但與女工的兒子們卻只分享四分之一的基因。因此，撫養自己的兒子而不是女工的兒子才符合女王的最大利益。 

雙方最大化其生殖成就導致王和工在貢獻不同雄性後代比例。女王喜歡生產自己的兒子，女工則喜歡生產自己的兒子，並且不會因為與侄子有良好的親緣關係而阻止姐妹產蛋。 

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.

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.

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.

提出的可能辨識機制


可能包括女王的碳氫化合物、女工的碳氫化合物或巢友識別。
女王的碳氫化合物

對弓背蟻屬 Camponotus floridanus的研究發現，其女王的卵表有特殊的碳氫化合物。在有女王產卵的群體，工蟻不產卵；而沒有女王產卵的群體，工蜂則繁殖。因此，碳氫化合物還可提醒工蟻停止繁殖。 目前還不清楚限制繁殖如何發生，或者減少工蜂繁殖所需的最低訊號閾值是多少。

女工的碳氫化合物

馬蟻的群體中，角質層化合物的存在引發了蟻群內部攻擊，特別是新舊繁殖工蟻之間。具體來說，老的生殖工蟻會在新雌性的觸角塗抹一種標記碳氫化合物，將它們識別為競爭對手和攻擊目標。

 同窩識別 

黑山蟻似乎還包括巢友識別。更多攻擊非同巢工蟻，並帶走其蛋。

 膜翅目例子 
趨同演化例子：

 蜜蜂 

最早發現的是西方蜜蜂（Apis mellifera）。普遍工蜂管制讓工蜂繁殖率極低（0.12%）。
蜜蜂透過吃來維持繁殖率（）。許多蜂巢工蜂卵巢活性減弱，產卵受抑制。 

據報道，小蜜蜂（Apis florea）也有行為。透過微衛星分析，沒有成熟的雄蜂具有非蜂王等位基因。因此，即使工蟻的卵巢已活化並且能夠產卵，工蟻的監管也確保了工蟻的功能性不孕。


據報導，東方蜜蜂雌性工蜂激活卵巢的比例高於義大利蜜蜂或小蜜蜂。當蜂后移除後，高達40%工蜂幾天內激活自己卵巢。然而，工蜂繼續吃工蟻的卵，顯示該物種的工者轄制並不涉及蜂后。

對蜜蜂屬來說也是類似。

勞者秩序也發生在原始的社會性熊蜂，例如欧洲熊蜂。

 螞蟻 

馬蟻 Pachycondyla inversa 會吃掉工蟻的卵（又稱），並攻擊產卵的蟻。

在 Gnamptogenys menadensis 的工蟻有時會交配產卵，而其它工蟻產用作食物的。由於其工蟻既可產雄性卵，也可產雌性卵，因此青睞勞者秩序。其勞者秩序可能有固定女工六肢（據報半數死亡），也可能拖出蟻群。 

Aphaenogaster smythiesi japonica 也顯示出勞者秩序的證據。當研究人員將重新組合有蟻后蟻群和無蟻后蟻群，工蟻會攻擊卵巢激活的工蟻。

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.

In the ponerine ant , 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.

 Wasps 
The tree wasp  also displays worker policing. Both egg-eating and aggression are reported in these wasp communities, and the queen also engages in the policing process. Similar behavior has been observed in the closely related species .

In colonies of the paper wasp, 中華馬蜂, 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  and monandrous colonies.

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.

In a test of worker policing, researchers examined the removal of worker-laid eggs in , 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.

The 黃邊胡蜂, 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.

 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.

 例外 

 無統治綜合症 

在極少數情況下，雌性工蜂的卵可以逃脫監管，這個過程被稱為無政府症候群。

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.

自私勞者秩序
, a 家蟻亞科 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 . 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.

This type of policy is common in  wasps where despite being more related to other workers' sons, worker-laid eggs were consumed by other workers.

 沒基因衝突的勞者秩序 
In the  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.

 參考 

 外部連結 
 Pachycondyla inversa 螞蟻維基的文章
 Gnamptogenys menadensis 螞蟻維基的文章
 Platythyrea punctata 螞蟻維基的文章
 Honeybee female worker egg-laying attempt  - video on YouTube
 Ant worker policing- video on YouTube
 Vespa crabro Worker policing in social hymenoptera

Category:行为生态学
Category:細腰亞目
Category:繁殖
Category:Evolutionary ecology