27The ongoing global temperature rise has led to increasing frequency of drought events, 28 negatively impacting vegetation and the living organisms relying on it. Extreme drought 29 killing host plants can clearly reduce herbivore fitness, but the impact of moderate host plant 30 water stress on insect herbivores can vary, and may even be beneficial. The Finnish Glanville 31 fritillary butterfly (Melitaea cinxia) has faced reduced precipitation in recent years, which has 32 impacted population dynamics. However, whether the negative effects depend on extreme 33 desiccation killing the host plant or moderate drought impacting plant quality remains unclear. 34 We assessed the performance of larvae fed on moderately water-stressed Plantago lanceolata 35 in terms of growth, gut microbial composition and immune response. We found that larvae 36 fed on water-stressed plants had better growth, a more heterogeneous bacterial community 37 and a shifted fungal community in the gut, and up-regulated the expression of one candidate 38 immune gene (pelle), whereas survival remained unaffected. Most of the measured traits 39 showed considerable variation due to family structure. Our data suggest that in temperate 40 regions moderate host plant water stress can positively shape resource acquisition of this 41 specialized insect herbivore, potentially by increasing nutrient accessibility or concentration.
42Potentially, the better larval performance may be mediated by a shift of the microbiota on 43 water-stressed plants, calling for further research especially on the understudied gut fungal 44 community. 45 46 Keywords: water stress, insect herbivore, gut microbiota, insect immunity 47 48 49 50 3 51 Introduction 52The present-day escalating greenhouse gas emissions have led to global climate change [1]. 53 One manifestation of this change is the increasing occurrence of events of drought or low 54 precipitation [1,2], impairing the biomass and species richness of plants, and consequently of 55 the herbivore species relying on them [3,4]. Insects can be differently affected by plant 56 drought, depending on the duration and frequency of the stress, and on the insect feeding 57 guild or species [5]. However, continuous water stress is generally expected to have a 58 negative impact [5]. Nonetheless, the degree of the water stress is also important, and extreme 59 desiccation killing the host plant is likely to have a negative impact on herbivore fitness.
60Importantly, insects should not be considered individually, but rather as holobionts composed 61 of the insect host and its microbial community [6]. Microbial communities play a role in 62 insect digestion, immunity and development, amongst others [7-11]. Microbiota composition 63 varies with host habitat and food source [12]. Contrarily to other insects, the impact and 64 spread of intracellular symbionts is quite limited in Lepidoptera [13], with recent work 65 suggesting that bacterial communities are often acquired through food and environment 66 [13,14]. There...