Developmental, morphological, and behavioural plasticity in the reproductive strategies of stink bugs and their egg parasitoids

L’environnement façonne la physiologie, la morphologie et le comportement des organismes par l’entremise de processus écologiques et évolutifs complexes et multidimensionnels. Le succès reproducteur des animaux est déterminé par la valeur adaptative d’un phénotype dans un environnement en modification constante selon une échelle temporelle d’une à plusieurs générations. De plus, les phénotypes sont façonnés par l’environnement, ce qui entraine des modifications adaptatives des stratégies de reproduction tout en imposant des contraintes. Dans cette thèse, considérant des punaises et leurs parasitoïdes comme organismes modèles, j’ai investigué comment plusieurs types de plasticité peuvent interagir pour influencer la valeur adaptative, et comment la plasticité des stratégies de reproduction répond à plusieurs composantes des changements environnementaux (qualité de l’hôte, radiation ultraviolette, température, invasion biologique). Premièrement, j’ai comparé la réponse comportementale et de traits d’histoire de vie à la variation de taille corporelle chez le parasitoïde Telenomus podisi Ashmead (Hymenoptera : Platygastridae), démontrant que les normes de réaction des comportements étaient plus souvent positives que celles des traits d’histoires de vie. Ensuite, j’ai démontré que la punaise prédatrice Podisus maculiventris Say (Hemiptera : Pentatomidae) peut contrôler la couleur de ses œufs, et que la pigmentation des œufs protège les embryons du rayonnement ultraviolet; une composante d’une stratégie complexe de ponte qui a évoluée en réponse à une multitude de facteurs environnementaux. Puis, j’ai testé comment le stress thermique affectait la dynamique de la mémoire du parasitoïde Trissolcus basalis (Wollaston) (Hymenoptera : Platygastridae) lors de l’apprentissage de la fiabilité des traces chimiques laissées par son hôte. Ces expériences ont révélé que des températures hautes et basses prévenaient l’oubli, affectant ainsi l’allocation du temps passé par les parasitoïdes dans des agrégats d’hôtes contenant des traces chimiques. J’ai aussi développé un cadre théorique général pour classifier les effets de la température sur l’ensemble des aspects comportementaux des ectothermes, distinguant les contraintes des adaptations. Finalement, j’ai testé l’habileté d’un parasitoïde indigène (T. podisi) à exploiter les œufs d’un nouveau ravageur invasif en agriculture, Halyomorpha halys Stål (Hemiptera : Pentatomidae). Les résultats ont montré que T. podisi attaque les œufs de H. halys, mais qu’il ne peut s’y développer, indiquant que le ravageur invasif s’avère un « piège évolutif » pour ce parasitoïde. Cela pourrait indirectement bénéficier aux espèces indigènes de punaises en agissant comme un puits écologique de ressources (œufs) et de temps pour le parasitoïde. Ces résultats ont des implications importantes sur la réponse des insectes, incluant ceux impliqués dans les programmes de lutte biologique, face aux changements environnementaux.The environment shapes the physiology, morphology, and behaviour of organisms through complex, multidimensional ecological and evolutionary processes. The reproductive success of individual animals is determined by how well their phenotype is suited to an environment that is constantly changing over single and multi-generational time scales. At the same time, phenotypes are shaped by the environment, which triggers adaptive modifications of animal reproductive strategies while also imposing important constraints. In this thesis, using stink bugs and their parasitoids as model organisms, I considered how several types of plasticity can interact to influence biological fitness, and how plasticity in reproductive strategies responds to several important components of environmental change (host quality, ultraviolet radiation, temperature, biological invasions). Firstly, I compared the response of behavioural and life history traits to body size variation in the parasitoid Telenomus podisi Ashmead (Hymenoptera: Platygastridae), finding that reaction norms of behavioural traits more often had positive slopes than life history traits. Next, I found that the predatory stink bug Podisus maculiventris Say (Hemiptera: Pentatomidae) can selectively control the colouration of its eggs. Egg pigmentation in this species protects embryos against ultraviolet radiation as part of a complex oviposition strategy that evolved in response to a suite of environmental factors. Then, I tested how thermal stress affects the memory dynamics of the parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Platygastridae) learning the reliability of chemical traces left by its host. These experiments revealed that both high and low stressful temperatures prevented forgetting, affecting the time allocation of parasitoids on patches of host chemical traces. I also developed a general framework to classify temperature’s effects on all aspects of ectotherm behaviour, distinguishing constraints from adaptive behavioural adjustments. Finally, I tested the ability of an indigenous parasitoid (T. podisi) to attack the eggs of a new invasive pest of agriculture, Halyomorpha halys Stål (Hemiptera: Pentatomidae). The results showed that T. podisi attacks the eggs of H. halys but cannot develop, demonstrating that the invasive pest is an “evolutionary trap” for indigenous parasitoids, which could indirectly benefit native stink bug species by acting as an egg and time sink for the parasitoid. These findings have important implications for how insects, including those involved in biological control programs, respond to environmental change

Thermal stress affects patch time allocation by preventing forgetting in a parasitoid wasp

Learning and memory allow animals to adjust their foraging strategies through experience. Despite the known impact of temperature on many aspects of the behavioral ecology of animals, memory retention in the face of realistic thermal stress has seldom been assessed. In the laboratory, we studied the behavioral expression of an egg parasitoid's (Trissolcus basalis) memory when exposed to thermal stress that could be encountered in nature. We hypothesized that thermal stress would disrupt memory consolidation and/or modify the optimality of memory retention, thus affecting patch time allocation strategies. Memory consolidation was resilient to 1h of thermal stress following an unrewarded experience (learning) on a patch of host-associated infochemicals. Neither high (40 °C) or low (10 °C) thermal stress changed the intensity of the experienced wasps' behavioral response relative to those held at a moderate temperature (25 °C). Next, we investigated how temperature stress could affect the parasitoids' memory retention ("forgetting"). When kept at a constant moderate temperature after learning, residence times of wasps retested on host cues increased relative to controls (naive wasps) over a period of 4 days as they presumably "forgot." However, both hot and cool daily temperature cycles prevented forgetting; the residence times of retested experienced wasps in these treatments did not change relative to controls over time. We discuss to what extent this may be an adaptive response by the parasitoids versus a physiological constraint imposed by temperature. Our findings contribute to an understanding of the impact of thermal stress on foraging strategies that involve learning and memory

Variation in levels of acceptance, developmental success, and abortion of Halyomorpha halys eggs by native North American parasitoids

Using native North American parasitoid species (Hymenoptera: Scelionidae) that often unsuccessfully attack the eggs of the invasive brown marmorated stink bug Halyomorpha halys (Hemiptera: Pentatomidae), we assessed variation in traits that may determine the parasitoids’ ability to adapt to the invasive host by either exploiting or avoiding H. halys eggs (acceptance, developmental success). We also assessed variation in the parasitoids’ ability to induce H. halys host egg abortion, which may contribute to biological control of H. halys in invaded areas. The first set of experiments evaluated intra- and interspecific variation using standardized laboratory tests with iso-female lines of Telenomus podisi and Trissolcus euschisti that included matching of detailed behavioural observations of acceptance with developmental outcomes. In a second set of experiments, we assessed how variation in developmental ability and abortion induction may affect levels of biological control by indigenous parasitoid species. We examined a broader sample of parasitoids that emerged from field collections of egg masses of an indigenous north American stink bug Podisus maculiventris in a region newly invaded by H. halys. Results from the first set of experiments showed high levels of acceptance of H. halys eggs among iso-female lines of parasitoids, but offspring development success was almost zero. H. halys egg abortion due to unsuccessful parasitism was often very low and varied among iso-female lines only for T. podisi. In the second set of experiments we never observed increases in abortion levels of Halyomorpha halys eggs above natural levels, even for the two species (T. euschisti and T. podisi) that were observed to oviposit in and abort H. halys eggs in the first set of experiments. We conclude that while there may be some variation in behavioural and physiological parameters mediating acceptance and abortion of H. halys eggs by native North American egg parasitoids, there does not appear to be significant variation in developmental success. Moreover, current biological control impact of H. halys eggs via host egg abortion is likely very low

Scientists' warning on climate change and insects

Climate warming is considered to be among the most serious of anthropogenic stresses to the environment, because it not only has direct effects on biodiversity, but it also exacerbates the harmful effects of other human-mediated threats. The associated consequences are potentially severe, particularly in terms of threats to species preservation, as well as in the preservation of an array of ecosystem services provided by biodiversity. Among the most affected groups of animals are insects—central components of many ecosystems—for which climate change has pervasive effects from individuals to communities. In this contribution to the scientists' warning series, we summarize the effect of the gradual global surface temperature increase on insects, in terms of physiology, behavior, phenology, distribution, and species interactions, as well as the effect of increased frequency and duration of extreme events such as hot and cold spells, fires, droughts, and floods on these parameters. We warn that, if no action is taken to better understand and reduce the action of climate change on insects, we will drastically reduce our ability to build a sustainable future based on healthy, functional ecosystems. We discuss perspectives on relevant ways to conserve insects in the face of climate change, and we offer several key recommendations on management approaches that can be adopted, on policies that should be pursued, and on the involvement of the general public in the protection effort.</p

Evidence of established brown marmorated stink bug populations in British Columbia, Canada

We report four new detections of invasive agricultural pest Halyomorpha halys (Stål) (Hempitera: Pentatomidae), the brown marmorated stink bug, in the Lower Mainland and Okanagan Valley regions of British Columbia (BC), Canada, in 2015 and 2016. These finds include two confirmed breeding populations, as well as homeowner collections at the same residence in two consecutive years. Preliminary comparisons of mitochondrial DNA haplotypes from these collections suggest that H. halys populations in BC are the result of movement and spread of existing populations in North America, likely from the Pacific Northwest USA

Switching to Tenofovir Alafenamide, Coformulated With Elvitegravir, Cobicistat, and Emtricitabine, in HIV-Infected Patients With Renal Impairment: 48-Week Results From a Single-Arm, Multicenter, Open-Label Phase 3 Study

BACKGROUND: Tenofovir alafenamide (TAF) is a novel tenofovir prodrug with improved renal and bone safety compared with TDF-containing regimens. We report the 48 week safety and efficacy of a once-daily single tablet regimen of elvitegravir 150 mg (E), cobicistat 150 mg (C), emtricitabine 200 mg (F), and TAF 10 mg (E/C/F/TAF) in HIV-1-infected patients with mild to moderate renal impairment. METHODS: We enrolled virologically suppressed HIV-1-infected subjects with estimated creatinine clearance (CrCl) 30-69 mL/min in a single-arm, open-label study to switch regimens to E/C/F/TAF. The primary endpoint was the change from baseline in glomerular filtration rate estimated using various formulae. This study is registered with ClinicalTrials.gov, number NCT01818596. FINDINGS: We enrolled and treated 242 patients with mean age 58 years, 18% Black, 39% hypertension, 14% diabetes. Through week 48, no significant change in estimated CrCl was observed. Two patients (0.8%) discontinued study drug for decreased creatinine clearance, neither had evidence of renal tubulopathy and both had uncontrolled hypertension. Subjects had significant improvements in proteinuria, albuminuria, and tubular proteinuria (P < 0.001 for all). Hip and spine bone mineral density significantly increased from baseline to week 48 (mean percent change +1.47 and +2.29, respectively, P < 0.05). Ninety-two percent (222 patients) maintained HIV-1 RNA <50 copies per milliliter at week 48. INTERPRETATION: Switch to E/C/F/TAF was associated with minimal change in GFR. Proteinuria, albuminuria and bone mineral density significantly improved. These data support the efficacy and safety of once daily E/C/F/TAF in HIV+ patients with mild or moderate renal impairment without dose adjustment

Flat photonic bands in guided modes of textured metallic microcavities

M. G. Salt and William L. Barnes, Physical Review B, Vol. 61, pp. 11125-11135 (2000). "Copyright © 2000 by the American Physical Society."A detailed experimental study of how wavelength-scale periodic texture modifies the dispersion of the guided modes of λ/2 metal-clad microcavities is presented. We first examine the case of a solid-state microcavity textured with a single, periodic corrugation. We explore how the depth of the corrugation and the waveguide thickness affect the width of the band gap produced in the dispersion of the guided modes by Bragg scattering off the periodic structure. We demonstrate that the majority of the corrugation depths studied dramatically modify the dispersion of the lowest-order cavity mode to produce a series of substantially flat bands. From measurements of how the central frequency of the band gap varies with direction of propagation of the guided modes, we determine a suitable two-dimensional texture profile for the production of a complete band gap in all directions of propagation. We then experimentally examine band gaps produced in the guided modes of such a two-dimensionally textured microcavity and demonstrate the existence of a complete band gap for all directions of propagation of the lowest-order TE-polarized mode. We compare our experimental results with those from a theoretical model and find good agreement. Implications of these results for emissive microcavity devices such as light-emitting diodes are discussed

Extreme genetic fragility of the HIV-1 capsid

Genetic robustness, or fragility, is defined as the ability, or lack thereof, of a biological entity to maintain function in the face of mutations. Viruses that replicate via RNA intermediates exhibit high mutation rates, and robustness should be particularly advantageous to them. The capsid (CA) domain of the HIV-1 Gag protein is under strong pressure to conserve functional roles in viral assembly, maturation, uncoating, and nuclear import. However, CA is also under strong immunological pressure to diversify. Therefore, it would be particularly advantageous for CA to evolve genetic robustness. To measure the genetic robustness of HIV-1 CA, we generated a library of single amino acid substitution mutants, encompassing almost half the residues in CA. Strikingly, we found HIV-1 CA to be the most genetically fragile protein that has been analyzed using such an approach, with 70% of mutations yielding replication-defective viruses. Although CA participates in several steps in HIV-1 replication, analysis of conditionally (temperature sensitive) and constitutively non-viable mutants revealed that the biological basis for its genetic fragility was primarily the need to coordinate the accurate and efficient assembly of mature virions. All mutations that exist in naturally occurring HIV-1 subtype B populations at a frequency &gt;3%, and were also present in the mutant library, had fitness levels that were &gt;40% of WT. However, a substantial fraction of mutations with high fitness did not occur in natural populations, suggesting another form of selection pressure limiting variation in vivo. Additionally, known protective CTL epitopes occurred preferentially in domains of the HIV-1 CA that were even more genetically fragile than HIV-1 CA as a whole. The extreme genetic fragility of HIV-1 CA may be one reason why cell-mediated immune responses to Gag correlate with better prognosis in HIV-1 infection, and suggests that CA is a good target for therapy and vaccination strategies

Global Peak in Atmospheric Radiocarbon Provides a Potential Definition for the Onset of the Anthropocene Epoch in 1965

Anthropogenic activity is now recognised as having profoundly and permanently altered the Earth system, suggesting we have entered a human-dominated geological epoch, the ‘Anthropocene’. To formally define the onset of the Anthropocene, a synchronous global signature within geological-forming materials is required. Here we report a series of precisely-dated tree-ring records from Campbell Island (Southern Ocean) that capture peak atmospheric radiocarbon (14C) resulting from Northern Hemisphere-dominated thermonuclear bomb tests during the 1950s and 1960s. The only alien tree on the island, a Sitka spruce (Picea sitchensis), allows us to seasonally-resolve Southern Hemisphere atmospheric 14C, demonstrating the ‘bomb peak’ in this remote and pristine location occurred in the last-quarter of 1965 (October-December), coincident with the broader changes associated with the post-World War II ‘Great Acceleration’ in industrial capacity and consumption. Our findings provide a precisely-resolved potential Global Stratotype Section and Point (GSSP) or ‘golden spike’, marking the onset of the Anthropocene Epoch

Scientists' warning on climate change and insects

Climate warming is considered to be among the most serious of anthropogenic stresses to the environment, because it not only has direct effects on biodiversity, but it also exacerbates the harmful effects of other human-mediated threats. The associated consequences are potentially severe, particularly in terms of threats to species preservation, as well as in the preservation of an array of ecosystem services provided by biodiversity. Among the most affected groups of animals are insects—central components of many ecosystems—for which climate change has pervasive effects from individuals to communities. In this contribution to the scientists' warning series, we summarize the effect of the gradual global surface temperature increase on insects, in terms of physiology, behavior, phenology, distribution, and species interactions, as well as the effect of increased frequency and duration of extreme events such as hot and cold spells, fires, droughts, and floods on these parameters. We warn that, if no action is taken to better understand and reduce the action of climate change on insects, we will drastically reduce our ability to build a sustainable future based on healthy, functional ecosystems. We discuss perspectives on relevant ways to conserve insects in the face of climate change, and we offer several key recommendations on management approaches that can be adopted, on policies that should be pursued, and on the involvement of the general public in the protection effort