Spider Mites Cause More Damage to Tomato in the Dark When Induced Defenses Are Lower.

Plants have evolved robust mechanisms to cope with incidental variation (e.g. herbivory) and periodical variation (e.g. light/darkness during the day-night cycle) in their environment. It has been shown that a plant's susceptibility to pathogens can vary during its day-night cycle. We demonstrated earlier that the spider mite Tetranychus urticae induces jasmonate- and salicylate-mediated defenses in tomato plants while the spider mite T. evansi suppresses these defenses probably by secreting salivary effector proteins. Here we compared induction/suppression of plant defenses; the expression of mite-effector genes and the amount of damage due to mite feeding during the day and during the night. T. urticae feeding upregulated the expression of jasmonate and salicylate marker-genes albeit significantly higher under light than under darkness. Some of these marker-genes were also upregulated by T. evansi-feeding albeit to much lower levels than by T. urticae-feeding. The expression of effector 28 was not affected by light or darkness in either mite species. However, the expression of effector 84 was considerably higher under light, especially for T. evansi. Finally, while T. evansi produced overall more feeding damage than T. urticae both mites produced consistently more damage during the dark phase than under light. Our results suggest that induced defenses are subject to diurnal variation possibly causing tomatoes to incur more damage due to mite-feeding during the dark phase. We speculate that mites, but especially T. evansi, may relax effector production during the dark phase because under these conditions the plant's ability to upregulate defenses is reduced.

Tetranychus evansi spider mite populations suppress tomato defenses to varying degrees.

Plant defense suppression is an offensive strategy of herbivores, in which they manipulate plant physiological processes to increase their performance. Paradoxically, defense suppression does not always benefit the defense-suppressing herbivores, because lowered plant defenses can also enhance the performance of competing herbivores and can expose herbivores to increased predation. Suppression of plant defense may therefore entail considerable ecological costs depending on the presence of competitors and natural enemies in a community. Hence, we hypothesize that the optimal magnitude of suppression differs among locations. To investigate this, we studied defense suppression across populations of Tetranychus evansi spider mites, a herbivore from South America that is an invasive pest of solanaceous plants including cultivated tomato, Solanum lycopersicum, in other parts of the world. We measured the level of expression of defense marker genes in tomato plants after infestation with mites from eleven different T. evansi populations. These populations were chosen across a range of native (South American) and non-native (other continents) environments and from different host plant species. We found significant variation at three out of four defense marker genes, demonstrating that T. evansi populations suppress jasmonic acid- and salicylic acid-dependent plant signaling pathways to varying degrees. While we found no indication that this variation in defense suppression was explained by differences in host plant species, invasive populations tended to suppress plant defense to a smaller extent than native populations. This may reflect either the genetic lineage of T. evansi-as all invasive populations we studied belong to one linage and both native populations to another-or the absence of specialized natural enemies in invasive T. evansi populations.

Distinct Signatures of Host Defense Suppression by Plant-Feeding Mites.

Tomato plants are attacked by diverse herbivorous arthropods, including by cell-content-feeding mites, such as the extreme generalist Tetranychus urticae and specialists like Tetranychus evansi and Aculops lycopersici. Mite feeding induces plant defense responses that reduce mite performance. However, T. evansi and A. lycopersici suppress plant defenses via poorly understood mechanisms and, consequently, maintain a high performance on tomato. On a shared host, T. urticae can be facilitated by either of the specialist mites, likely due to the suppression of plant defenses. To better understand defense suppression and indirect plant-mediated interactions between herbivorous mites, we used gene-expression microarrays to analyze the transcriptomic changes in tomato after attack by either a single mite species (T. urticae, T. evansi, A. lycopersici) or two species simultaneously (T. urticae plus T. evansi or T. urticae plus A. lycopersici). Additionally, we assessed mite-induced changes in defense-associated phytohormones using LC-MS/MS. Compared to non-infested controls, jasmonates (JAs) and salicylate (SA) accumulated to higher amounts upon all mite-infestation treatments, but the response was attenuated after single infestations with defense-suppressors. Strikingly, whereas 8 to 10% of tomato genes were differentially expressed upon single infestations with T. urticae or A. lycopersici, respectively, only 0.1% was altered in T. evansi-infested plants. Transcriptome analysis of dual-infested leaves revealed that A. lycopersici primarily suppressed T. urticae-induced JA defenses, while T. evansi dampened T. urticae-triggered host responses on a transcriptome-wide scale. The latter suggests that T. evansi not solely down-regulates plant gene expression, but rather directs it back towards housekeeping levels. Our results provide valuable new insights into the mechanisms underlying host defense suppression and the plant-mediated facilitation of competing herbivores.

Inbreeding depression in a critically endangered carnivore.

Harmful effects arising from matings between relatives (inbreeding) is a long-standing observation that is well founded in theory. Empirical evidence for inbreeding depression in natural populations is however rare because of the challenges of assembling pedigrees supplemented with fitness traits. We examined the occurrence of inbreeding and subsequent inbreeding depression using a unique data set containing a genetically verified pedigree with individual fitness traits for a critically endangered arctic fox (Vulpes lagopus) population. The study covered nine years and was comprised of 33 litters with a total of 205 individuals. We recorded that the present population was founded by only five individuals. Over the study period, the population exhibited a tenfold increase in average inbreeding coefficient with a final level corresponding to half-sib matings. Inbreeding mainly occurred between cousins, but we also observed two cases of full-sib matings. The pedigree data demonstrated clear evidence of inbreeding depression on traditional fitness traits where inbred individuals displayed reduced survival and reproduction. Fitness traits were however differently affected by the fluctuating resource abundande. Inbred individuals born at low-quality years displayed reduced first-year survival, while inbred individuals born at high-quality years were less likely to reproduce. The documentation of inbreeding depression in fundamental fitness traits suggests that inbreeding depression can limit population recovery. Introducing new genetic material to promote a genetic rescue effect may thus be necessary for population long-term persistence.

Life history traits in a cyclic ecosystem: a field experiment on the arctic fox.

The reproduction of many species depends strongly on variation in food availability. The main prey of the arctic fox in Fennoscandia are cyclic small rodents, and its number of litters and litter size vary depending on the phase of the rodent cycle. In this experiment, we studied if the arctic fox adjusts its reproduction as a direct response to food abundance, in accordance with the food limitation hypothesis, or if there are additional phase-dependent trade-offs that influence its reproduction. We analysed the weaning success, i.e. proportion of arctic fox pairs established during mating that wean a litter in summer, of 422 pairs of which 361 were supplementary winter fed, as well as the weaned litter size of 203 litters of which 115 were supplementary winter fed. Females without supplementary winter food over-produced cubs in relation to food abundance in the small rodent increase phase, i.e. the litter size was equal to that in the peak phase when food was more abundant. The litter size for unfed females was 6.38 in the increase phase, 7.11 in the peak phase and 3.84 in the decrease phase. The litter size for supplementary winter-fed litters was 7.95 in the increase phase, 10.61 in the peak phase and 7.86 in the decrease phase. Thus, feeding had a positive effect on litter size, but it did not diminish the strong impact of the small rodent phase, supporting phase-dependent trade-offs in addition to food determining arctic fox reproduction.

Pearsonema (syn Capillaria) plica associated cystitis in a Fennoscandian arctic fox (Vulpes lagopus: a case report.

The bladderworm Pearsonema (syn Capillaria) plica affects domestic dogs and wild carnivores worldwide. A high prevalence in red foxes (Vulpes vulpes) has been reported in many European countries. P. plica inhabits the lower urinary tract and is considered to be of low pathogenic significance in dogs mostly causing asymptomatic infections. However, a higher level of pathogenicity has been reported in foxes. A severe cystitis associated with numerous bladderworms was found in a captive arctic fox (Vulpes lagopus) originating from the endangered Fennoscandian arctic fox population. To our knowledge this is the first description of P. plica infection in an arctic fox.