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社会性昆虫的任务分配与划分

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Task allocation and partitioning is the way that tasks are chosen, assigned, subdivided, and coordinated within a colony of social insects. Task allocation and partitioning gives rise to the division of labor often observed in social insect colonies, whereby individuals specialize on different tasks within the colony (e.g., "foragers", "nurses"). Communication is closely related to the ability to allocate tasks among individuals within a group.

Task allocation and partitioning is the way that tasks are chosen, assigned, subdivided, and coordinated within a colony of social insects. Task allocation and partitioning gives rise to the division of labor often observed in social insect colonies, whereby individuals specialize on different tasks within the colony (e.g., "foragers", "nurses"). Communication is closely related to the ability to allocate tasks among individuals within a group.

任务分配和划分是在一群社会性昆虫中，任务被选择、分配、细分和协调的方式。任务分配和分配引起社会昆虫群落中经常观察到的分工，即个体在群落中专注于不同的任务(如“采集者”、“护士”)。沟通与团队中个人分配任务的能力密切相关。

This entry focuses exclusively on social insects. For information on human task allocation and partitioning, see division of labour, task analysis, and workflow.

This entry focuses exclusively on social insects. For information on human task allocation and partitioning, see division of labour, task analysis, and workflow.

这个条目专门关注社会性昆虫。有关人工任务分配和分区的信息，请参见分工、任务分析和工作流。


Definitions

  • Task allocation "... is the process that results in specific workers being engaged in specific tasks, in numbers appropriate to the current situation. [It] operates without any central or hierarchical control..."[1] The concept of task allocation is individual-centric. It focuses on decisions by individuals about what task to perform. However, different biomathematical models give different weights to inter-individual interactions vs. environmental stimuli.[1]
  • Task partitioning is the division of one task into sequential actions done by more than one individual.[2] The focus here is on the task, and its division, rather than on the individuals performing it. For example, "hygienic behavior" is a task in which worker bees uncap and remove diseased brood cells that may be affected by American foulbrood (Paenibacillus larvae) or the parasitic mite Varroa destructor.[3] In this case, individual bees often focus on either uncapping or removing diseased brood. Therefore, the task is partitioned, and performed by multiple individuals.[4]


Introduction

Social living provides a multitude of advantages to its practitioners, including predation risk reduction, environmental buffering, food procurement, and possible mating advantages. The most advanced form of sociality is eusociality, characterized by overlapping generations, cooperative care of the young, and reproductive division of labor, which includes sterility or near-sterility of the overwhelming majority of colony members. With few exceptions, all the practitioners of eusociality are insects of the orders Hymenoptera (ants, bees, and wasps), Isoptera (termites), Thysanoptera (thrips), and Hemiptera (aphids).

社会生活为其从业者提供了许多优势，包括减少捕食风险、环境缓冲、获取食物和可能的交配优势。社会性的最高级形式是真社会性、拥有属性重叠的世代、合作照顾年轻人以及繁殖分工，其中包括绝大多数群体成员的不育或近不育。除了极少数的例外，所有真正意义上的实践者都是膜翅目(蚂蚁、蜜蜂和黄蜂)、等翅目(白蚁)、胸腺翅目(蓟马)和半翅目(蚜虫)的昆虫。

Social living provides a multitude of advantages to its practitioners, including predation risk reduction, environmental buffering, food procurement, and possible mating advantages. The most advanced form of sociality is eusociality, characterized by overlapping generations, cooperative care of the young, and reproductive division of labor, which includes sterility or near-sterility of the overwhelming majority of colony members. With few exceptions, all the practitioners of eusociality are insects of the orders Hymenoptera (ants, bees, and wasps), Isoptera (termites), Thysanoptera (thrips), and Hemiptera (aphids).[5][6]

Social insects have been extraordinarily successful ecologically and evolutionarily. This success has at its most pronounced produced colonies 1) having a persistence many times the lifespan of most individuals of the colony, and 2) numbering thousands or even millions of individuals.

从生态学和进化学的角度来看，社会性昆虫是非常成功的。这种成功在其最显著的产生殖民地1)具有多倍于殖民地大多数个体寿命的持久性，2)数以千计甚至数百万计的个体。

Social insects have been extraordinarily successful ecologically and evolutionarily. This success has at its most pronounced produced colonies 1) having a persistence many times the lifespan of most individuals of the colony, and 2) numbering thousands or even millions of individuals.

Social insects can exhibit division of labor with respect to non-reproductive tasks, in addition to the aforementioned reproductive one. In some cases this takes the form of markedly different, alternative morphological development (polymorphism), as in the case of soldier castes in ants, termites, thrips, and aphids, while in other cases it is age-based (temporal polyethism), as with honey bee foragers, who are the oldest members of the colony (with the exception of the queen). Evolutionary biologists are still debating the fitness-advantage gained by social insects due to their advanced division of labor and task allocation, but hypotheses include: increased resilience against a fluctuating environment, reduced energy costs of continuously switching tasks, increased longevity of the colony as a whole, or reduced rate of pathogen transmission.

除了上述的繁殖任务之外，社会性昆虫在非繁殖任务方面也表现出分工。在某些情况下，这种现象表现为明显不同的替代形态发育(多态性) ，如蚂蚁、白蚁、蓟马和蚜虫中的士兵种姓，而在其他情况下，这种现象是基于年龄的(暂时性) ，如蜜蜂觅食者，他们是群体中年龄最大的成员(除了蜂王)。进化生物学家仍在争论社会性昆虫由于其高级的劳动分工和任务分配而获得的适应优势，但假设包括: 对波动环境的适应能力增强，连续切换任务的能量成本降低，群体整体寿命延长，或者水平传播减少。

Social insects can exhibit division of labor with respect to non-reproductive tasks, in addition to the aforementioned reproductive one. In some cases this takes the form of markedly different, alternative morphological development (polymorphism), as in the case of soldier castes in ants, termites, thrips, and aphids, while in other cases it is age-based (temporal polyethism), as with honey bee foragers, who are the oldest members of the colony (with the exception of the queen). Evolutionary biologists are still debating the fitness-advantage gained by social insects due to their advanced division of labor and task allocation, but hypotheses include: increased resilience against a fluctuating environment, reduced energy costs of continuously switching tasks, increased longevity of the colony as a whole, or reduced rate of pathogen transmission.[7][8]

Division of labor, large colony sizes, temporally-changing colony needs, and the value of adaptability and efficiency under Darwinian competition, all form a theoretical basis favoring the existence of evolved communication in social insects. Beyond the rationale, there is well-documented empirical evidence of communication related to tasks; examples include the waggle dance of honey bee foragers, trail marking by ant foragers such as the red harvester ants, and the propagation via pheromones of an alarm state in Africanized honey bees.

分工、大群体规模、时变群体需求以及达尔文竞争下的适应性和效率价值，构成了社会昆虫进化交流存在的理论基础。除了基本原理之外，还有很多与任务相关的沟通经验证明，例如蜜蜂采集蜂的摇摆舞，蚂蚁采集蜂的踪迹标记，以及非洲化蜜蜂通过信息素的报警状态传播[7][8]。

Division of labor, large colony sizes, temporally-changing colony needs, and the value of adaptability and efficiency under Darwinian competition, all form a theoretical basis favoring the existence of evolved communication in social insects.[9][10][11] Beyond the rationale, there is well-documented empirical evidence of communication related to tasks; examples include the waggle dance of honey bee foragers, trail marking by ant foragers such as the red harvester ants, and the propagation via pheromones of an alarm state in Africanized honey bees.


Worker Polymorphism

One of the most well known mechanisms of task allocation is worker polymorphism, where workers within a colony have morphological differences. This difference in size is determined by the amount of food workers are fed as larvae, and is set once workers emerge from their pupae. Workers may vary just in size (monomorphism) or size and bodily proportions (allometry). An excellent example of the monomorphism is in bumblebees (Bombus spp.). Bumblebee workers display a large amount of body size variation which is normally distributed. The largest workers may be ten times the mass of the smallest workers. Worker size is correlated with several tasks: larger workers tend to forage, while smaller workers tend to perform brood care and nest thermoregulation. Size also affects task efficiency. Larger workers are better at learning, have better vision, carry more weight, and fly at a greater range of temperatures. However, smaller workers are more resistant to starvation. In other eusocial insects as well, worker size can determine what polymorphic role they become. For instance, larger workers in Myrmecocystus mexicanus (a North America species of honeypot ant) tend to become repletes, or workers so engorged with food that they become immobile and act a living food storage for the rest of the colonies.

最为人熟知的任务分配机制之一是工作者多态性，即群体内的工作者存在形态学上的差异。这种大小的差异取决于工蚁幼虫喂食的食物量，并且是在工蚁从蛹中出来后确定的。工人可能只是大小不同(单形)或大小和身体比例(异速生长)。单态的一个很好的例子是熊蜂(Bombus spp.)。大黄蜂工蜂体型变异较大，呈正态分布。最大的工人可能是最小的工人的十倍。工蚁的大小与几个任务有关: 较大的工蚁倾向于觅食，而较小的工蚁倾向于照顾幼仔和体温调节。大小也会影响任务的效率。体型较大的工作者学习能力更强，视力更好，携带更多的重量，在更大的温度范围内飞行。然而，体型较小的工人对饥饿的抵抗力更强。在其他真社会昆虫以及，工人大小可以决定什么多态性的作用，他们成为。例如，墨西哥蚁属(Myrmecocystus mexicanus)(一种北美洲的蜜罐蚂蚁)体型较大的工蚁往往会饱食一顿，或者工蚁饱食一顿，变得行动不便，成为其余殖民地的活食物储存库。

One of the most well known mechanisms of task allocation is worker polymorphism, where workers within a colony have morphological differences. This difference in size is determined by the amount of food workers are fed as larvae, and is set once workers emerge from their pupae. Workers may vary just in size (monomorphism) or size and bodily proportions (allometry). An excellent example of the monomorphism is in bumblebees (Bombus spp.). Bumblebee workers display a large amount of body size variation which is normally distributed. The largest workers may be ten times the mass of the smallest workers. Worker size is correlated with several tasks: larger workers tend to forage, while smaller workers tend to perform brood care and nest thermoregulation. Size also affects task efficiency. Larger workers are better at learning, have better vision, carry more weight, and fly at a greater range of temperatures. However, smaller workers are more resistant to starvation.[12] In other eusocial insects as well, worker size can determine what polymorphic role they become. For instance, larger workers in Myrmecocystus mexicanus (a North America species of honeypot ant) tend to become repletes, or workers so engorged with food that they become immobile and act a living food storage for the rest of the colonies.[13]


In many ants and termites, on the other hand, workers vary in both size and bodily proportions, which have a bimodal distribution. This is present in approximately one in six ant genera. In most of these there are two developmentally distinct pathways, or castes, into which workers can develop. Typically members of the smaller caste are called minors and members of the larger caste are called majors or soldiers. There is often variation in size within each caste. The term soldiers may be apt, as in Cephalotes, but in many species members of the larger caste act primarily as foragers or food processors. In a few ant species, such as certain Pheidole species, there is a third caste, called supersoldiers.

另一方面，在许多蚂蚁和白蚁中，工蚁在体型和身体比例上都有所不同，它们有一个双峰分布。这种现象大约存在于六个蚂蚁属中的一个。在大多数情况下，有两种不同的发育路径，或者说种姓，工人们可以进入这两种发育路径。通常较小种姓的成员被称为未成年人，较大种姓的成员被称为少校或士兵。每个种姓的大小往往不同。士兵这个词可能比较贴切，比如在 Cephalotes，但在许多物种中，较大种姓的成员主要扮演着觅食者或食物加工者的角色。在一些蚂蚁物种中，比如某些大头蚁物种，存在第三种等级，称为超级蚂蚁。

In many ants and termites, on the other hand, workers vary in both size and bodily proportions, which have a bimodal distribution. This is present in approximately one in six ant genera. In most of these there are two developmentally distinct pathways, or castes, into which workers can develop. Typically members of the smaller caste are called minors and members of the larger caste are called majors or soldiers. There is often variation in size within each caste. The term soldiers may be apt, as in Cephalotes, but in many species members of the larger caste act primarily as foragers or food processors. In a few ant species, such as certain Pheidole species, there is a third caste, called supersoldiers.


Temporal polyethism

Temporal polyethism is a mechanism of task allocation, and is ubiquitous among eusocial insect colonies. Tasks in a colony are allocated among workers based on their age. Newly emerged workers perform tasks within the nest, such as brood care and nest maintenance, and progress to tasks outside the nest, such as foraging, nest defense, and corpse removal as they age. In honeybees, the youngest workers exclusively clean cells, which is then followed by tasks related to brood care and nest maintenance from about 2–11 days of age. From 11– 20 days, they transition to receiving and storing food from foragers, and at about 20 days workers begin to forage. Similar temporal polyethism patterns can be seen in primitive species of wasps, such as Ropalidia marginata as well as the eusocial wasp Vespula germanica. Young workers feed larvae, and then transition to nest building tasks, followed by foraging. Many species of ants also display this pattern. This pattern is not rigid, though. Workers of certain ages have strong tendencies to perform certain tasks, but may perform other tasks if there is enough need. For instance, removing young workers from the nest will cause foragers, especially younger foragers, to revert to tasks such as caring for brood. These changes in task preference are caused by epigenetic changes over the life of the individual. Honeybee workers of different ages show substantial differences in DNA methylation, which causes differences in gene expression. Reverting foragers to nurses by removing younger workers causes changes in DNA methylation similar to younger workers.

时间聚合现象是一种任务分配机制，在群落昆虫中普遍存在。群体中的任务是根据工人的年龄分配的。新出现的工蚁在巢内执行任务，例如照料幼鸟和维护巢穴，并且在巢外执行任务，例如觅食，筑巢防御，以及随着年龄增长移除尸体。在蜜蜂中，最年轻的工蜂专门清洁蜂房，然后从2-11天开始接下来的任务是照顾幼蜂和维护蜂巢。从11-20天开始，它们从采集者那里接收和储存食物，大约20天后，工蚁开始觅食。类似的时间聚合模式可以在原始种类的黄蜂中看到，比如边缘黄蜂和德国黄胡蜂。年轻的工蚁喂养幼虫，然后过渡到筑巢任务，接着是觅食。许多种类的蚂蚁也表现出这种模式。不过，这种模式并不僵化。某些年龄段的工人有强烈的倾向去完成某些任务，但是如果有足够的需要，他们也可以完成其他任务。例如，把年轻的工蚁赶出巢穴会导致觅食蚁，特别是年轻的觅食蚁，回到照顾幼蚁的工作上来。这些任务偏好的变化是由个体生命中的表观遗传变化引起的。不同年龄的蜜蜂工人在 DNA 甲基化方面存在显著差异，这导致了基因表达的差异。通过清除年轻工人的方式将采集者还原给护士，会导致与年轻工人相似的 DNA 甲基化改变。

Temporal polyethism is a mechanism of task allocation, and is ubiquitous among eusocial insect colonies. Tasks in a colony are allocated among workers based on their age. Newly emerged workers perform tasks within the nest, such as brood care and nest maintenance, and progress to tasks outside the nest, such as foraging, nest defense, and corpse removal as they age. In honeybees, the youngest workers exclusively clean cells, which is then followed by tasks related to brood care and nest maintenance from about 2–11 days of age. From 11– 20 days, they transition to receiving and storing food from foragers, and at about 20 days workers begin to forage.[14] Similar temporal polyethism patterns can be seen in primitive species of wasps, such as Ropalidia marginata as well as the eusocial wasp Vespula germanica. Young workers feed larvae, and then transition to nest building tasks, followed by foraging.[15] Many species of ants also display this pattern.[16] This pattern is not rigid, though. Workers of certain ages have strong tendencies to perform certain tasks, but may perform other tasks if there is enough need. For instance, removing young workers from the nest will cause foragers, especially younger foragers, to revert to tasks such as caring for brood.[17] These changes in task preference are caused by epigenetic changes over the life of the individual. Honeybee workers of different ages show substantial differences in DNA methylation, which causes differences in gene expression. Reverting foragers to nurses by removing younger workers causes changes in DNA methylation similar to younger workers.[18]

Temporal polyethism is not adaptive because of maximized efficiency; indeed older workers are actually more efficient at brood care than younger workers in some ant species.

暂时的多元性并不是因为最大化的效率而具有适应性; 事实上，在一些蚂蚁种类中，年长的工蚁实际上比年轻的工蚁更有效率地照顾幼蚁。

Temporal polyethism is not adaptive because of maximized efficiency; indeed older workers are actually more efficient at brood care than younger workers in some ant species.[17] Rather it allows workers with the lowest remaining life expectancy to perform the most dangerous tasks. Older workers tend to perform riskier tasks, such as foraging, which has high risks of predation and parasitism, while younger workers perform less dangerous tasks, such as brood care. If workers experience injuries, which shortens their life expectancies, they will start foraging sooner than healthy workers of the same age.[19]


Response-Threshold Model

A dominant theory of explaining the self-organized division of labor in social insect societies such as honey bee colonies is the Response-Threshold Model. It predicts that individual worker bees have inherent thresholds to stimuli associated with different tasks. Individuals with the lowest thresholds will preferentially perform that task. The Response-Threshold Model only provides for effective task allocation in the honey bee colony if thresholds are varied among individual workers. This variation originates from the considerable genetic diversity among worker daughters of a colony due to the queen’s multiple matings.

响应阈值模型是解释社会性昆虫群体(如蜜蜂群体)自组织分工的一个主要理论。它预测，个别工蜂对与不同任务相关的刺激有固有的阈值。最低阈值的个体将优先执行这项任务。响应阈值模型只提供了有效的任务分配在蜜蜂群体中，如果阈值是不同的工人。这种变异源于蚁后多次交配所导致的蚁群工蜂女儿之间相当大的遗传多样性。

A dominant theory of explaining the self-organized division of labor in social insect societies such as honey bee colonies is the Response-Threshold Model. It predicts that individual worker bees have inherent thresholds to stimuli associated with different tasks. Individuals with the lowest thresholds will preferentially perform that task.[7] Stimuli could include the “search time” that elapses while a foraging bee waits to unload her nectar and pollen to a receiver bee at the hive, the smell of diseased brood cells, or any other combination of environmental inputs that an individual worker bee encounters.[20] The Response-Threshold Model only provides for effective task allocation in the honey bee colony if thresholds are varied among individual workers. This variation originates from the considerable genetic diversity among worker daughters of a colony due to the queen’s multiple matings.[21]


Network representation of information flow and task allocation

To explain how colony-level complexity arises from the interactions of several autonomous individuals, a network-based approach has emerged as a promising area of social insect research. Social insect colonies can be viewed as a self-organized network, in which interacting elements (i.e. nodes) communicate with each other. As decentralized networks, colonies are capable of distributing information rapidly which facilitates robust responsiveness to their dynamic environments. The efficiency of information flow is critical for colony-level flexibility because worker behavior is not controlled by a centralized leader but rather is based on local information.

为了解释群落层面的复杂性是如何从几个自治个体的相互作用中产生的，基于网络的方法已经成为社会昆虫研究的一个有前途的领域。社会性昆虫群落可以看作是一个自组织网络，其中的相互作用元素(即昆虫群落中的相互作用元素)是昆虫群落的一个重要组成部分。节点)相互通信。作为分散网络，殖民地能够快速分布信息，这有利于对其动态环境的鲁棒响应。信息流的效率对群体层次的灵活性至关重要，因为员工的行为不受集中式领导的控制，而是基于局部信息。

To explain how colony-level complexity arises from the interactions of several autonomous individuals, a network-based approach has emerged as a promising area of social insect research. Social insect colonies can be viewed as a self-organized network, in which interacting elements (i.e. nodes) communicate with each other. As decentralized networks, colonies are capable of distributing information rapidly which facilitates robust responsiveness to their dynamic environments.[22] The efficiency of information flow is critical for colony-level flexibility because worker behavior is not controlled by a centralized leader but rather is based on local information.


Social insect networks are often non-randomly distributed, wherein a few individuals act as ‘hubs,’ having disproportionately more connections to other nestmates than other workers in the colony. Computer simulations of this particular interaction network demonstrated that inter-individual variation in connectivity patterns expedites information flow among nestmates.

社会昆虫网络通常是非随机分布的，其中一些个体充当“中心” ，它们与其他同伴的联系比群体中的其他工蚁要多得多。计算机模拟这种特殊的互动网络表明，个体之间连接模式的差异加速了同伴之间的信息流动。

Social insect networks are often non-randomly distributed, wherein a few individuals act as ‘hubs,’ having disproportionately more connections to other nestmates than other workers in the colony.[22] In harvester ants, the total interactions per ant during recruitment for outside work is right-skewed, meaning that some ants are more highly connected than others.[23] Computer simulations of this particular interaction network demonstrated that inter-individual variation in connectivity patterns expedites information flow among nestmates.


Task allocation within a social insect colony can be modeled using a network-based approach, in which workers are represented by nodes, which are connected by edges that signify inter-node interactions. Workers performing a common task form highly connected clusters, with weaker links across tasks. These weaker, cross-task connections are important for allowing task-switching to occur between clusters. In Formica fusca L. ant colonies, a network analysis of spatial effects on feeding and the regulation of food storage revealed that food is distributed heterogeneously within colony, wherein heavily loaded workers are located centrally within the nest and those storing less food were located at the periphery.

社会昆虫群体内的任务分配可以用基于网络的方法来建模，其中工蚁由表示节点的边连接起来，表示节点间的相互作用。执行共同任务的工作人员形成了高度连接的集群，各任务之间的链接较弱。这些较弱的跨任务连接对于允许在集群之间进行任务切换非常重要。在福米加褐孔蚁群体中，对取食的空间效应和食物储存调节的网络分析表明，食物在群体中的分布是不均匀的，重负荷的工蚁集中在巢内，而储存较少食物的工蚁集中在周围。

Task allocation within a social insect colony can be modeled using a network-based approach, in which workers are represented by nodes, which are connected by edges that signify inter-node interactions. Workers performing a common task form highly connected clusters, with weaker links across tasks. These weaker, cross-task connections are important for allowing task-switching to occur between clusters.[22] This approach is potentially problematic because connections between workers are not permanent, and some information is broadcast globally, e.g. through pheromones, and therefore does not rely on interaction networks. One alternative approach to avoid this pitfall is to treat tasks as nodes and workers as fluid connections.


Studies of inter-nest pheromone trail networks maintained by super-colonies of Argentine ants (Linepithema humile) have shown that different colonies establish networks with very similar topologies. Insights from these analyses revealed that these networks – which are used to guide workers transporting brood, workers and food between nests – are formed through a pruning process, in which individual ants initially create a complex network of trails, which are then refined to eliminate extraneous edges, resulting in a shorter, more efficient inter-nest network.

由阿根廷蚂蚁的超级群体维持的巢间信息素追踪网络的研究表明，不同的群体建立的网络具有非常相似的拓扑结构。从这些分析中得出的结论是，这些网络是通过一个修剪过程形成的，在这个过程中，蚂蚁个体最初创建了一个复杂的路径网络，然后再进一步细化以消除无关的边缘，从而形成一个更短、更有效的巢间网络。

To demonstrate how time and space constraints of individual-level interactions affect colony function, social insect network approaches can also incorporate spatiotemporal dynamics. These effects can impose upper bounds to information flow rate in the network. For example, the rate of information flow through Temnothorax rugatulus ant colonies is slower than would be predicted if time spent traveling and location within the nest were not considered.[24] In Formica fusca L. ant colonies, a network analysis of spatial effects on feeding and the regulation of food storage revealed that food is distributed heterogeneously within colony, wherein heavily loaded workers are located centrally within the nest and those storing less food were located at the periphery.[25]


Long-term stability of interaction networks has been demonstrated in Odontomachus hastatus ants, in which initially highly connected ants remain highly connected over an extended time period. Conversely, Temnothorax rugatulus ant workers are not persistent in their interactive role, which might suggest that social organization is regulated differently among different eusocial species. Traditionally, workers are the nodes of the graph, but Fewell prefers to make the tasks the nodes, with workers as the links. O'Donnell has coined the term "worker connectivity" to stand for "communicative interactions that link a colony's workers in a social network and affect task performance". See also the review of task partitioning by Ratnieks and Anderson. Because of the often limited quantities and limited precision of data from which to calculate parameters values in non-human behavior studies, such models should generally be kept simple. Therefore, we generally should not expect models for social insect task allocation or task partitioning to be as elaborate as human workflow ones, for example.

长期稳定的相互作用网络已经在[ http://www.antwiki.org/wiki/odontomachus_hastatus ]蚂蚁身上得到了证明，在这种蚂蚁身上，最初高度连接的蚂蚁在较长的时间内保持高度连接。相反，[ http://www.antwiki.org/wiki/temnothorax_rugatulus ]蚂蚁工蚁并不坚持它们的互动角色，这可能表明不同群居物种之间的社会组织是不同的。传统上，工作者是图中的节点，但 Fewell 更喜欢将任务作为节点，工作者作为链接。O’ donnell 创造了“工作者连接”这个术语来代表“沟通交互作用，这种交互作用将一个群体的工作者联系在一个社会网络中，并影响工作绩效”。请参阅 Ratnieks 和 Anderson 对任务分区的评论。由于在非人类行为研究中计算参数值的数据往往数量有限，精度也有限，因此这些模型通常应该保持简单。因此，我们通常不应该期望社会昆虫任务分配或任务划分模型像人类工作流模型那样精细。

Studies of inter-nest pheromone trail networks maintained by super-colonies of Argentine ants (Linepithema humile) have shown that different colonies establish networks with very similar topologies.[26] Insights from these analyses revealed that these networks – which are used to guide workers transporting brood, workers and food between nests – are formed through a pruning process, in which individual ants initially create a complex network of trails, which are then refined to eliminate extraneous edges, resulting in a shorter, more efficient inter-nest network.


Long-term stability of interaction networks has been demonstrated in Odontomachus hastatus ants, in which initially highly connected ants remain highly connected over an extended time period.[27] Conversely, Temnothorax rugatulus ant workers are not persistent in their interactive role, which might suggest that social organization is regulated differently among different eusocial species.[24]

With increased data, more elaborate metrics for division of labor within the colony become possible. Gorelick and Bertram survey the applicability of metrics taken from a wide range of other fields. They argue that a single output statistic is desirable, to permit comparisons across different population sizes and different numbers of tasks. But they also argue that the input to the function should be a matrix representation (of time spent by each individual on each task), in order to provide the function with better data. They conclude that "... normalized matrix-input generalizations of Shannon's and Simpson's index ... should be the indices of choice when one wants to simultaneously examine division of labor amongst all individuals in a population".

随着数据的增加，更详细的分工指标成为可能。Gorelick 和 Bertram 调查了从其他领域广泛采集的指标的适用性。他们认为，单一的输出统计数据是可取的，以允许在不同的人口规模和不同的任务数量之间进行比较。但是他们也认为，为了给函数提供更好的数据，函数的输入应该是一个矩阵表示(每个人在每个任务上花费的时间)。他们得出结论: “ ... ... 香农和辛普森指数的矩阵输入归纳法... ... 应该成为一个人想要同时检验一个人群中所有个体之间的分工时的选择指数。”。


Note that these indexes, used as metrics of biodiversity, now find a place measuring division of labor.

请注意，这些指数，作为生物多样性的衡量标准，现在找到了一个衡量劳动分工的地方。

A network is pictorially represented as a graph, but can equivalently be represented as an adjacency list or adjacency matrix.[28] Traditionally, workers are the nodes of the graph, but Fewell prefers to make the tasks the nodes, with workers as the links.[29][30] O'Donnell has coined the term "worker connectivity" to stand for "communicative interactions that link a colony's workers in a social network and affect task performance".[30] He has pointed out that connectivity provides three adaptive advantages compared to individual direct perception of needs:[30]

  1. It increases both the physical and temporal reach of information. With connectivity, information can travel farther and faster, and additionally can persist longer, including both direct persistence (i.e. through pheromones), memory effects, and by initiating a sequence of events.
  1. It can help overcome task inertia and burnout, and push workers into performing hazardous tasks. For reasons of indirect fitness, this latter stimulus should not be necessary if all workers in the colony are highly related genetically, but that is not always the case.
  1. Key individuals may possess superior knowledge, or have catalytic roles. Examples, respectively, are a sentry who has detected an intruder, or the colony queen.

O'Donnell provides a comprehensive survey, with examples, of factors that have a large bearing on worker connectivity.[30] They include:

  • size of the interacting group, especially if the network has a modular structure
  • sender distribution (i.e. a small number of controllers vs. numerous senders)
  • strength of the interaction effect, which includes strength of the signal sent, recipient sensitivity, and signal persistence (i.e. pheromone signal vs. sound waves)
  • recipient memory, and its decay function
  • socially-transmitted inhibitory signals, as not all interactions provide positive stimulus
  • specificity of both the signal and recipient response
  • signal and sensory modalities, and activity and interaction rates


Task taxonomy and complexity

Category:Behavioral ecology

类别: 行为生态学

Anderson, Franks, and McShea have broken down insect tasks (and subtasks) into a hierarchical taxonomy; their focus is on task partitioning and its complexity implications. They classify tasks as individual, group, team, or partitioned; classification of a task depends on whether there are multiple vs. individual workers, whether there is division of labor, and whether subtasks are done concurrently or sequentially. Note that in their classification, in order for an action to be considered a task, it must contribute positively to inclusive fitness; if it must be combined with other actions to achieve that goal, it is considered to be a subtask. In their simple model, they award 1, 2, or 3 points to the different tasks and subtasks, depending on its above classification. Summing all tasks and subtasks point values down through all levels of nesting allows any task to be given a score that roughly ranks relative complexity of actions.[31] See also the review of task partitioning by Ratnieks and Anderson.[2]

Category:Superorganisms

类别: 超级有机体


Category:Insect behavior

类别: 昆虫行为


This page was moved from wikipedia:en:Task allocation and partitioning of social insects. Its edit history can be viewed at 社会性昆虫的任务分配与划分/edithistory

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  2. ↑ 2.0 2.1 Francis L. W. Ratnieks & Carl Anderson (1999). "Task partitioning in insect societies". Insectes Sociaux. 47 (2): 95–108. doi:10.1007/s000400050119. S2CID 24435668.
  3. ↑ Arathi, H. S.; Burns, I.; Spivak, M. (2000). "Ethology of hygienic behaviour in the honey bee Apis mellifera L-(Hymenoptera : Apidae): Behavioural repertoire of hygienic bees". Ethology. 106 (4): 365–379. doi:10.1046/j.1439-0310.2000.00556.x.
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  5. ↑ John R. Krebs; Nicholas B. Davies (1987). An Introduction to Behavioural Ecology (2nd ed.). Blackwell Scientific Publications. p. 291. 
  6. ↑ Ross H. Crozier; Pekka Pamilo (1996). "Introduction". Evolution of Social Insect Colonies. Sex Allocation and Kin Selection. Oxford Series in Ecology and Evolution. Oxford University Press. pp. 4–8. ISBN 978-0-19-854942-0. 
  7. ↑ 7.0 7.1 7.2 Duarte, A.; Pen, I.; Keller, L.; Weissing, F. J. (2012). "Evolution of self-organized division of labor in a response threshold model". Behavioral Ecology and Sociobiology. 66 (6): 947–957. doi:10.1007/s00265-012-1343-2. PMC 3353103. PMID 22661824.
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  10. ↑ Sasha R. X. Dall, Luc-Alain Giraldeau, Ola Olsson, John M. McNamara & David W. Stephens (2005). "Information and its use by animals in evolutionary ecology" (PDF). Trends in Ecology & Evolution. 20 (4): 187–193. doi:10.1016/j.tree.2005.01.010. PMID 16701367.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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  13. ↑ Rissing, Steven W (1984). "Replete Caste Production and Allometry of Workers in the Honey Ant, Myrmecocystus mexicanus Wesmael (Hymenoptera: Formicidae)". Journal of the Kansas Entomological Society. 57 (2): 347–350.
  14. ↑ Thomas D. Seeley (1982). "Adaptive significance of the age polyethism schedule in honeybee colonies". Behavioral Ecology and Sociobiology. 11 (4): 287–293. doi:10.1007/BF00299306. JSTOR 4599548. S2CID 37953822.
  15. ↑ Dhruba Naug & Raghavendra Gadagkar (1998). "The role of age in temporal polyethism in a primitively eusocial wasp" (PDF). Behavioral Ecology and Sociobiology. 42 (1): 37–47. doi:10.1007/s002650050409. JSTOR 4601416. S2CID 14615486.
  16. ↑ Bert Hölldobler & E. O. Wilson (1990). The Ants. Cambridge, MA: Harvard University Press. ISBN 978-0-674-04075-5. 
  17. ↑ 17.0 17.1 Muscedere, ML; Willey TA; Traniello JFA (2009). "Age and task efficiency in the ant Pheidole dentata: young minor workers are not specialist nurses". Animal Behaviour. 77 (4): 911–918. doi:10.1016/j.anbehav.2008.12.018. S2CID 53161750.
  18. ↑ Herb, BR; Wolschin F; Hansen K; Aryee MJ; Langmead B; Irizarry R; Amdam GV; AP Feinbert (2012). "Reversible switching between epigenetic states in honeybee behavioral subcastes". Nature Neuroscience. 15 (10): 1371–1375. doi:10.1038/nn.3218. PMC 3518384. PMID 22983211.
  19. ↑ Kuszewska, K; Woyciechowski M (2012). "Reversion in honeybee, Apis mellifera, workers with different life expectancies". Animal Behaviour. 85: 247–253. doi:10.1016/j.anbehav.2012.10.033. S2CID 54346742.
  20. ↑ Thenius, R.; Schmickl, T.; Crailsheim, K. (2008). "Optimisation of a honeybee-colony's energetics via social learning based on queuing delays". Connection Science. 20 (2–3): 193–210. doi:10.1080/09540090802091982. S2CID 27954392.
  21. ↑ Tarapore, D.; Floreano, D.; Keller, L. (2010). "Task-dependent influence of genetic architecture and mating frequency on division of labour in social insect societies". Behavioral Ecology and Sociobiology. 64 (4): 675–684. doi:10.1007/s00265-009-0885-4. S2CID 6084107.
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  23. ↑ "The effect of individual variation on the structure and function of interaction networks in harvester ants". 171.66.127.193. 2011-11-07. Retrieved 2013-11-26.
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  25. ↑ "Collective regulatory stock management and spatiotemporal dynamics of the food flow in ants". The FASEB Journal. 2012-07-01. Retrieved 2013-11-26.
  26. ↑ "Structure and formation of ant transportation networks". Journal of the Royal Society Interface. 2011-09-07. Retrieved 2013-11-26.
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  28. ↑ Michael Goodrich & Roberto Tamassia (2002). Algorithm Design. Wiley. p. 296. ISBN 978-0-471-38365-9. 
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  30. ↑ 30.0 30.1 30.2 30.3 S. O'Donnell & S. J. Bulova (2007). "Worker connectivity: a review of the design of worker communication systems and their effects on task performance in insect societies". Insectes Sociaux. 54 (3): 203–210. doi:10.1007/s00040-007-0945-6. S2CID 10839817.
  31. ↑ Carl Anderson, Nigel R. Franks & Daniel W. McShea (2001). "The complexity and hierarchical structure of tasks in insect societies". Animal Behaviour. 62 (4): 643–651. doi:10.1006/anbe.2001.1795. S2CID 7551221.
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