Ecological theory and observational evidence suggest that symbiotic interactions such as cleaning symbioses can shift from mutualism to parasitism. However, field experimental evidence documenting these shifts has never been reported for a cleaning symbiosis. Here, we demonstrate shifts in a freshwater cleaning symbiosis in a system involving crayfish and branchiobdellid annelids. Branchiobdellids have been shown to benefit their hosts under some conditions by cleaning material from host crayfish's gill filaments. The system is uniquely suited as an experimental model for symbiosis due to ease of manipulation and ubiquity of the organisms. In three field experiments, we manipulated densities of worms on host crayfish and measured host growth in field enclosures. In all cases, the experiments revealed shifts from mutualism to parasitism: host crayfish growth was highest at intermediate densities of branchiobdellid symbionts, while high symbiont densities led to growth that was lower or not significantly different from 0-worm controls. Growth responses were consistent even though the three experiments involved different crayfish and worm species and were performed at different locations. Results also closely conformed to a previous laboratory experiment using the same system. The mechanism for these shifts appears to be that branchiobdellids switched from cleaning host gills at intermediate densities of worms to consuming host gill tissue at high densities. These outcomes clearly demonstrate shifts along a symbiosis continuum with the maximum benefits to the host at intermediate symbiont densities. At high symbiont densities, benefits to the host disappear, and there is some evidence for a weak parasitism. These are the first field experimental results to demonstrate such shifts in a cleaning symbiosis.
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Visualizing thermoregulation (endothermy vs. ectothermy) in animals can be challenging for students in undergraduate biology courses. Data-driven, hands-on laboratory activities can enhance student learning while reinforcing application of the scientific method. This article describes a visual-learning, inquiry-based activity that can be applied to introductory high school and college biology laboratories with the use of easily accessible technology (a smartphone and an inexpensive thermal camera attachment). Students generate hypotheses and qualitatively observe real-time thermal images of live endothermic and ectothermic animals. This activity is effective in engaging students by allowing them to visualize thermoregulation and body temperature. We provide suggestions for modifying the activity to further investigate animal behavior related to temperature regulation.
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