Multiple stressors: using the honeybee model BEEHAVE to explore how spatial and temporal forage stress affects colony resilience
Multiple stressors: using the honeybee model BEEHAVE to explore how spatial and temporal forage stress affects colony resilience
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多重压力源：使用蜜蜂模型 BEEHAVE 探索时空草料压力如何影响蜂群恢复力
作者：
Horn J
过去十年观察到的蜜蜂蜜蜂群体死亡率增加的原因尚不清楚。由于迄今为止单因解释的证据尚不明确，因此通常假设涉及多种压力源。我们在这里关注草料可用性的各个方面，与其他压力因素相比，这些方面受到的关注较少，因为实际上不可能凭经验来探索它们。我们将蜂群模型 BEEHAVE（将蜂巢内动态和觅食联系起来）应用到程式化的景观环境中，以探索觅食距离、饲料供应和“饲料间隙”（即蜜蜂找不到任何花蜜和花粉的时期）如何影响蜂群恢复能力及其背后的机制。我们发现，群体灭绝主要是由觅食距离驱动的，但觅食间隙的时间对灭绝时间的影响最强。即使草料供应足以维持生存，6 月或 7 月对 15 天草料缺口的敏感性最高。草料供应通过对蚁后产卵率的级联效应、发育成成年工蚁的新育雏阶段的减少、花粉债务、缺乏护理劳动力以及减少觅食活动来影响蚁群。七月的饲料缺口导致产卵量减少，育雏期死亡率增加，而此时蜂王的季节性产卵率达到最高，随着时间的推移，导致蜂群失败。我们的结果表明，不合时宜的饲料缺口与整体饲料供应不足相互作用，会降低蜂群的恢复力。原则上使蜂群能够应对特定景观和年份中不同饲料供应（例如产蛋量的减少）的现有调节机制仅具有一定的能力。我们的结果是假设的，因为它们是从简化的景观设置中获得的，但它们与现有的经验知识一致。它们提供了充足的机会在对照实验中测试草料胁迫的预测影响。
The causes underlying the increased mortality of honeybeeApis melliferacolonies observed over the past decade remain unclear. Since so far the evidence for monocausal explanations is equivocal, involvement of multiple stressors is generally assumed. We here focus on various aspects of forage availability, which have received less attention than other stressors because it is virtually impossible to explore them empirically. We applied the colony model BEEHAVE, which links within‐hive dynamics and foraging, to stylized landscape settings to explore how foraging distance, forage supply, and “forage gaps”, i.e. periods in which honeybees cannot find any nectar and pollen, affect colony resilience and the mechanisms behind. We found that colony extinction was mainly driven by foraging distance, but the timing of forage gaps had strongest effects on time to extinction. Sensitivity to forage gaps of 15 days was highest in June or July even if otherwise forage availability was sufficient to survive. Forage availability affected colonies via cascading effects on queen's egg‐laying rate, reduction of new‐emerging brood stages developing into adult workers, pollen debt, lack of workforce for nursing, and reduced foraging activity. Forage gaps in July led to reduction in egg‐laying and increased mortality of brood stages at a time when the queen's seasonal egg‐laying rate is at its maximum, leading to colony failure over time. Our results demonstrate that badly timed forage gaps interacting with poor overall forage supply reduce honeybee colony resilience. Existing regulation mechanisms which in principle enable colonies to cope with varying forage supply in a given landscape and year, such as a reduction in egg‐laying, have only a certain capacity. Our results are hypothetical, as they are obtained from simplified landscape settings, but they are consistent with existing empirical knowledge. They offer ample opportunities for testing the predicted effects of forage stress in controlled experiments.
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