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The stress-gradient hypothesis (SGH) predicts that the strength and frequency of facilitative interactions increase monotonically with increasing environmental stress, but some empirical studies have found this decrease at extreme stress levels, suggesting a hump-shaped SGH instead. However, empirical studies of the SGH are often hindered by confounding resource and non-resource stress gradients. Nepenthes pitcher plants trap animal prey using modified-leaf pitfall traps which are also inhabited by organisms known as inquilines. Inquilines may assist pitchers in the digestion of trapped prey. This interaction is known as a digestive mutualism and is both mutualistic and facilitative by definition. Inquiline species may also facilitate each other via processing chain commensalisms. We used in vitro experiments to examine the isolated effect of resource stress on the outcomes of two facilitative interactions: (i) digestive mutualism—facilitation of pitcher nutrient sequestration by two inquiline dipteran larvae, culicids and phorids; (ii) processing chain commensalism—facilitation between these two inquiline taxa. The net nutritional benefit of phorids on N. gracilis was found to conform more to a monotonic rather than hump-shaped SGH model. However, the effect of culicids on N. gracilis and the effects of culicids and phorids on each other were weak. These findings provide compelling evidence that changes in facilitation along an isolated resource stress gradient conform to the predictions of the monotonic SGH model rather than that of the revised hump-shaped model, and highlight the importance of isolating stress gradients in empirical tests of the SGH. [ABSTRACT FROM AUTHOR]

• Processing chain commensalism (PCC) is a type of resource conversion interaction. • Long-term outcomes of PCCs are invariant to the specification of functional responses. • SGH in PCCs requires leakage of the downstream resource to occur. • These findings are likely to apply to other resource conversion-type interactions. The stress-gradient hypothesis (SGH) in ecology predicts that the strength and frequency of positive interspecific interactions, including processing chain commensalisms (PCCs), increase with environmental stress. Although observed in some empirical PCC studies, a recent theoretical study of PCCs using a consumer–resource-type model found that, given the model's assumptions, the SGH never occurs. To investigate if this is a true reflection of PCCs or merely an artefact of the model, in this study, we modified this earlier model formulation by incorporating generalized, monotonically increasing resource uptake functions in place of linear functions, and added a term to represent the spontaneous leakage of the downstream resource to the environment. Mathematical analyses of the model revealed two key insights: 1) the specific algebraic forms of the functional responses of the species in a PCC do not affect the long-term behaviour of the system; 2) the SGH can occur in a facilitative interaction only if the consumer-independent leakage rate of the downstream resource exceeds the consumer-independent input rate. The first insight shows that the outcomes of consumer–resource interactions are robust to details of the functional responses when the functional responses are monotonically increasing, while the second insight shows that the SGH is not a universal feature of positive interactions but instead holds only under a well-defined set of conditions which may vary between PCC interactions and the environmental contexts in which they take place. [ABSTRACT FROM AUTHOR]