The formation of the nestmate recognition template in ants and its effect on behavioural variation

我们对他人的行为取决于我们是否认识他们，例如通过他们的脸。我们将感知到的面孔与存储在记忆中的面孔原型——模板——进行比较。虽然识别的其他方面（生产成分，如提示表达；行动成分，如决策规则）得到了很好的研究，但以模板为基本方面的感知成分却没有得到很好的研究。目前，无论是人类还是动物，我们对模板中存储的信息以及模板是如何形成的都知之甚少。群居昆虫是研究模板形成的一个特别好的模型，因为工蜂对入侵者的攻击性不同，我们表明感知成分是造成一些差异的原因。巢友识别模板的传统观点将其描述为“平均群体气味”的代表，这对所有的巢友都是相同的。然而，模板可能部分是自我参照的，例如，个体自身气味和平均群体气味的组合（“新模板”）。这可以解释已经观察到的行为差异：模板，以及识别，会随着每个人自己的气味而变化。对于模板的形成，也有两种可供选择的假设：模板可能是由习惯形成的：蚂蚁“习惯”了自己的群体气味。或者，联想学习可能发挥作用：蚂蚁可能将群体气味与与其他蚂蚁互动时的好或坏经历联系起来，如果个体经历不同的互动，这可能会导致个体间的差异。如果蚂蚁学会将非配偶的气味与负面经历（例如打架）联系起来，这将特别明显，这将提高对来自熟悉的非配偶群体的入侵者的检测，只有那些以前遇到过非配偶群体的个体。我们从“讨厌的邻居效应”的实验中获得了后者的初步证据，即对非巢伴侣的熟悉会增加蚂蚁对特定非巢伴侣群体的攻击性。为了测试自参照模板或模板的联想学习是否对观察到的巢识别变化负责，我们将操纵工蚁相对于其群体气味的气味，并操纵个体经历的相互作用。我们也将直接阻碍联想学习与药理学干预。最后，我们将分析大脑和触角的基因表达，以跟踪我们的实验操作引起的分子效应。然后，我们将这些影响与不同工蜂形态之间的分子变异进行比较，这些工蜂形态的攻击性各不相同。我们的项目将阐明长期存在且目前备受争议的问题，即巢识别模板是如何形成的，这对我们理解行为变化和一般识别过程具有重要意义。

Our behaviour towards others depends on whether we recognise them, for example by their faces. We compare the perceived face with prototypes of faces stored in our memory: the templates. While other aspects of recognition (production component, e.g. cue expression; action component, e.g. decision rules) are well studied, the perception component, of which template is a fundamental aspect, is not. Currently, we have little understanding from any system, human or animal, of the information stored in templates, and how templates are formed. Social insects are a particularly good model to study template formation because workers differ in their aggression towards intruders and we showed that the perception component is responsible for some of the variation.Traditional views of the nestmate recognition template describe it as a representation of the “average colony odour”, which is identical for all nestmates. However, the template could be partly self-referent, e.g. a combination of an individual's own odour and the average colony odour (“Newey template”). This would explain the behavioural variation that has been observed: Templates, and thus recognition, would vary with each individual's own odour. For template formation, there are two alternative hypotheses as well: The template could be formed by habituation: Ants “get used” to their own colony odour. Alternatively, associative learning could play a role: Ants may associate colony odours to good or bad experiences in interactions with other ants, which is likely to cause inter-individual variation if individuals experience different interactions. This would be particularly evident if ants learn to associate non-nestmate odours with negative experiences (e.g. fights), which would improve the detection of intruders from familiar non-nestmate colonies only in those individuals that previously encountered the non-nestmate colony in question. We have preliminary evidence for the latter from experiments on the nasty neighbour effect, where familiarity with non-nestmates increases an ant's aggression towards the specific non-nestmate colony.To test whether self-referent templates or associative learning of templates are responsible for the observed variation in nestmate recognition, we will manipulate the odours of ant workers relative to their colony odour and manipulate the interactions experienced by individuals. We will also directly impede associative learning with pharmacological interventions. Finally, we will analyse brain and antennal gene expression to track molecular effects caused by our experimental manipulations. We will then compare these effects to the molecular variation among different worker morphs, which vary in their aggression. Our project will shed light on the long-standing and currently hotly debated question of how nestmate recognition templates are formed, with implications for our understanding behavioural variation and recognition processes in general.