Integrative neuroimmunomodulation of gastrointestinal function during enteric parasitism.

Enteric helminths have a significant impact on the structure, function, and neural control of the gastrointestinal (GI) tract of the host. Interactions between the host's nervous and immune systems redirect activity in neuronal circuits intrinsic to the gut into an alternative repertoire of defensive and adaptive motor programs. Gut inflammation and activation of the enteric neuroimmune axis play integral roles in the dynamic interaction between host and parasite that occurs at the mucosal surface. Three inter-related themes are stressed in this review to underscore the pivotal role that neural control mechanisms play in the host's GI tract functional responses to enteric parasitism. First, we address the discovery that signaling molecules of both parasite and host origin can reorient the dynamic ecology of enteric host-parasite interactions. Second, we explore what has been learned from investigations of altered gut propulsive and secretomotor reflex activities that occur during enteric parasitic infections and the emerging picture derived from these studies that elucidates how nerves help facilitate and orchestrate functional reorganization of the parasitized gut. Third, we provide an overview of the direct impact that enteric parasitism has on nerve cell function and neurotransmission pathways in both the enteric and central nervous systems of the host. In summary, this review highlights and clarifies the complex mechanisms underlying integrative neuroimmunophysiological responses to the presence of both invasive and noninvasive enteric helminths and identifies directions for future research investigations in this highly important but understudied area.     

Ecological and physiological adaptations in insect parasitism

Ecological and physiological adaptations of the tachinid parasite Lydella grisescens R-D, to its hosts were studied experimentally. The factors affecting host finding and larviposition of the adult females were evaluated. The frass of four lepidopterous larvae, including two natural hosts, the European corn borer Ostrinia nubilalis (Hbn.) and the stalk borer Papaipema nebris (Guen.), and two non-hosts, the armyworm Pseudaletia unipuncta (Haworth) and the cat-tail borer Archanata oblonga Grt., was found to attract the Lydella females and stimulate them to larviposit. The attracting material in frass is soluble in ethanol and water, and seems to be a metabolic product of the caterpillars rather than a product of plant fermentation.Artificial parasitization by implantation of Lydella larvae in the host caterpillars was employed to determine the physiological suitability of various natural and unnatural hosts to the parasite. Along with the two natural hosts, the European corn borer and the stalk borer, an unnatural host, the greater wax moth Galleria mellonella L. was found to be a suitable host. Artificial parasitization was not successful with the armyworm, the imported cabbageworm Pieris rapae L. and the yellow mealworm Tenebrio molitor L. Lydella larvae were able to penetrate the cuticle of Galleria larvae when they were present together. The biological significance and practical implication of these findings are discussed.

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Zusammenfassung Die ökologischen und physiologischen Anpassungen der parasitischen Tachinide Lydella grisescens R.-D. an ihre Wirte wurden experimentell untersucht. Die Faktoren, welche die Wirtsfindung und das Larvenabsetzen der erwachsenen Weibchen betreffen, wurden ermittelt. Der Kot von vier Schmetterlingslarven, nämlich zweier natürlicher Wirte: des Maiszünslers Ostrinia nubilalis (Hbn.) und des Stengelbohrers Papaipema nebris (Guen.), und zweier Nichtwirte; des Heerwurmes Pseudaletia unipuncta (Haworth) und des Katzenschwanz-Bohrers Archanata oblonga Grt. erwies sich als anziehend für die Lydella-Weibchen und veranlaßte sie zum Absetzen von Larven. Das anlockende Material im Kot ist in Alkohol und Wasser löslich und scheint eher ein Stoffwechselprodukt der Raupen als ein Zersetzungsprodukt der Pflanze zu sein.Künstliche Parasitierung durch Implantation von Lydella-Larven in Wirtsraupen wurde benutzt, um die physiologische Eignung der verschiedenen natürlichen und unnatürlichen Wirte für den Parasiten zu bestimmen. Neben den zwei natürlichen Wirten, dem Maiszünsler und dem Stengelbohrer, wurde ein unnatürlicher Wirt, die Große Wachsmotte, Galleria mellonella L., als geeigneter Wirt nachgewiesen. Die künstliche Parasitierung blieb dagegen beim Heerwurm, dem (eingeschleppten) Kohlweißling Pieris rapae L. und dem Mehlwurm Tenebrio molitor L. erfolglos. Lydella-Larven sind fähig, die Kutikula der Galleria-Larven zu durchdringen. Die biologische und praktische Bedeutung dieser Befunde wird diskutiert.

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Paper No. 5673 Scientific Journal Series, Minnesota Experiment Station, St. Paul, Minnesota, 55101.     

Levels of herbivory and parasitism in host hybrid zones

Hybrid zones often are intermediate in their habitat, and have individuals that are intermediate genetically to those of either parental species. Recently, a number of studies has addressed the distribution of herbivores and parasites in hybrid zones of their hosts. Mechanisms that underlie these distributions include behavioral, environmental and genotypic differences between hybrids and their parental species.     

寄主大小及寄生顺序对蝇蛹佣小蜂寄生策略的影响

寄主大小模型认为寄生蜂后代性比与寄主大小相关,寄生蜂倾向于在大寄主上产出更多雌性后代,在小寄主上产出更多雄性后代.探讨了以家蝇蛹为寄主时,蝇蛹佣小蜂后代产量和性比变化;单次寄生情况下,寄主大小及寄生顺序对寄生蜂后代性比等影响.结果表明,蝇蛹佣小蜂的产卵期为(8.93±3.34)d,单头雌蜂能产雌性后代(34.11±16.34)头和雄性后代(11.04±8.87)头,且雄性百分比为0.24±0.11.随成蜂日龄的增大,寄生蜂产生雄性后代的比率显著增加.蝇蛹佣小蜂在寄生家蝇蛹时,会优先选择寄生个体较大的蛹;在单次寄生的情况下,蝇蛹佣小蜂倾向于在较大的家蝇蛹内产出更多的雌性后代.     

Brood parasitism selects for no defence in a cuckoo host

In coevolutionary arms races, like between cuckoos and their hosts, it is easy to understand why the host is under selection favouring anti-parasitism behaviour, such as egg rejection, which can lead to parasites evolving remarkable adaptations to ‘trick’ their host, such as mimetic eggs. But what about cases where the cuckoo egg is not mimetic and where the host does not act against it? Classically, such apparently non-adaptive behaviour is put down to evolutionary lag: given enough time, egg mimicry and parasite avoidance strategies will evolve. An alternative is that absence of egg mimicry and of anti-parasite behaviour is stable. Such stability is at first sight highly paradoxical. I show, using both field and experimental data to parametrize a simulation model, that the absence of defence behaviour by Cape bulbuls (Pycnonotus capensis) against parasitic eggs of the Jacobin cuckoo (Clamator jacobinus) is optimal behaviour. The cuckoo has evolved massive eggs (double the size of bulbul eggs) with thick shells, making it very hard or impossible for the host to eject the cuckoo egg. The host could still avoid brood parasitism by nest desertion. However, higher predation and parasitism risks later in the season makes desertion more costly than accepting the cuckoo egg, a strategy aided by the fact that many cuckoo eggs are incorrectly timed, so do not hatch in time and hence do not reduce host fitness to zero. Selection will therefore prevent the continuation of any coevolutionary arms race. Non-mimetic eggs and absence of defence strategies against cuckoo eggs will be the stable, if at first sight paradoxical, result.     

Field evidence of host size-dependent parasitism in two manipulative parasites

The distribution of parasites within host natural populations has often been found to be host age-dependent. Host mortality induced by parasites is the commonest hypothesis proposed for explaining this pattern. Despite its potential importance in ecology, the parasitism intensity in relation with the host age has rarely been studied in the field. The 2 manipulative acanthocephalans, Polymorphus minutus and Pomphorhynchus laevis, use the amphipod Gammarus pulex as an intermediate host, and their infection intensity and incidence among G. pulex populations were examined by analyzing 2 large samples of hosts collected in eastern France. Both parasites had low prevalence in the host populations, but their mean abundances were highly related with gammarid age. For the 2 acanthocephalans, results reported a disappearance or an absence of heavily infected hosts in the older host age classes. These results suggested that parasites that alter intermediate host behavior for enhancing their transmission success to the definitive host reduce the survival of their intermediate host. In conclusion, manipulative parasites might act as a mechanism regulating the density of gammarid populations.     

Persistence of host defence behaviour in the absence of avian brood parasitism

The fate of host defensive behaviour in the absence of selection from brood parasitism is critical to long-term host-parasite coevolution. We investigated whether New World Bohemian waxwings Bombycilla garrulus that are allopatric from brown-headed cowbird Molothrus ater and common cuckoo Cuculus canorus parasitism have retained egg rejection behaviour. We found that egg rejection was expressed by 100 per cent of Bohemian waxwings. Our phylogeny revealed that Bohemian and Japanese waxwings Bombycilla japonica were sister taxa, and this clade was sister to the cedar waxwing Bombycilla cedrorum. In addition, there was support for a split between Old and New World Bohemian waxwings. Our molecular clock estimates suggest that egg rejection may have been retained for 2.8-3.0 Myr since New World Bohemian waxwings inherited it from their common ancestor with the rejecter cedar waxwings. These results support the 'single trajectory' model of host-brood parasite coevolution that once hosts evolve defences, they are retained, forcing parasites to become more specialized over time.     

Proteomic interactions between the parasitoid Diachasmimorpha longicaudata and the oriental fruit fly,Bactrocera dorsalis during host parasitism

The oriental fruit fly, Bactrocera dorsalis, is an important agricultural pest and biological control is one of the most effective control methodologies. We conducted an investigation on the molecular response of the fruit fly to parasitism by the larval parasitoid, Diachasmimorpha longicaudata using two-dimensional gel electrophoresis and mass spectroscopy. We identified 285 differentially expressed protein spots (109 proteins) during parasitism. The molecular processes affected by parasitism varied at different time point during development. Transferrin and muscle specific protein 20 are the only two proteins differentially expressed that play a role in host immunity 24 h after parasitism. Developmental and metabolic proteins from parasitoids (transferrin and enolase) were up-regulated to ensure establishment and early development of parasitoids 48 h post parasitism. 72 h after parasitism, larval cuticle proteins, transferrin and CREG1 were overexpressed to support the survival of parasitoids while host metabolism proteins and parasitoid regulatory proteins were down-regulated. Host development slowed down while parasitoid development went up at 96 h after parasitism. All developmental, regulatory, structural, and metabolic proteins were expressed at their optimum at 120 h post parasitism. Host development was reduced, metabolism and regulatory proteins were strongly involved in the activities. The development deteriorated further at 144 h after parasitism. Enolase and CREG1 were indicators of parasitoid survival. Hexamerin and transferrin from the parasitoid was peaked at 168–216 h after parasitism, strongly indicating that parasitoid would survive. This study represents the first report that reveals the molecular players involved in the interaction between the host and parasitoid.     

Predation risk, host immune response, and parasitism

Predation risk may affect the allocation priorities of limitingresources by potential prey. Investment in immune function shouldreceive reduced priority, when hosts are exposed to predatorsbecause of the costs of immune function. We tested this hypothesisby randomly exposing adult house sparrows, Passer domesticus,to either a cat, Felis catus, or a rabbit, Oryctolagus cuniculus,for 6 h while assessing their ability to raise a T-cell–mediatedimmune response to a challenge with phytohemagglutinin. Sparrowsexposed to a cat had a significant reduction of, on average,18% and 36% in T-cell response in two different experimentscompared with sparrows that were exposed to a rabbit. In a fieldexperiment with a barn owl, Tyto alba, or a rock dove, Columbalivia, placed next to a nest-box during laying, we found a meanreduction in T-cell–mediated immune response of 20%. Inmales, the reduction in cell-mediated immune response owingto cat exposure increased with increasing size of the badge,which is a secondary sexual character, but only during the breedingseason. In a third experiment, house sparrows were either exposedto a barn own, T. alba, or a rock dove, C. livia, and developmentof malarial infections was recorded during the following 6 weeks.Individual sparrows exposed to a predator had a higher prevalenceand intensity of Haemoproteus malarial infection than did controlindividuals. Therefore, exposure to predators reduced theirability of hosts to cope with parasitism mediated through effectson immune function.     

Mating and host density affect host feeding and parasitism in two species of whitefly parasitoids

Abstract The parasitoids in the genera of Encarsia and Eretmocerus (Hymenoptera: Aphelinidae) are important biological control agents of whiteflies, and some of them not only parasitize hosts but also kill them with strong host‐feeding capacity. Two whitefly parasitoid species, Encarsia sophia and Eretmocerus melanoscutus were examined to determine if mating and host density affected their host feeding and parasitism. The whitefly host, Bemisia tabaci, was presented to these two wasp species in densities of 10, 20, 30, 40, 50 and 60 third‐instar nymphs per clip cage. Mated whitefly parasitoid females fed on more hosts than unmated females under a range of host densities (under all six host densities for En. sophia; under the densities of 40 nymphs or more for Er. melanoscutus). Meanwhile, mated females parasitized more whitefly nymphs than unmated females under all host densities for both species. With increase of host density, mated or unmated Er. melanoscutus females killed more hosts by host feeding and parasitism. Mated En. sophia females killed more hosts by host feeding with increase of host density, whereas unmated females did not parasitze whitefly nymphs at all. Our results suggest that only mated female parasitoids with host‐feeding behavior should be released in crop systems to increase their bio‐control efficiency.     

How parasitism and pollution affect the physiological homeostasis of aquatic hosts

Parasitism poses a serious threat to hosts under certain circumstances, while the well-being of organisms is also negatively affected by environmental pollution. Little information is available on the simultaneous effects of parasites and pollutants on the physiological homeostasis of organisms. The present paper demonstrates that parasites: (i) may influence the metabolism of pollutants in infected hosts, (ii) interact with pollution in synergistic or antagonistic ways, and (iii) may induce physiological reactions in hosts which were thought to be pollutant-induced. Experimental studies on the uptake and accumulation of metals by fish reveal that fish infected with acanthocephalans have lower metal levels than uninfected hosts; e.g. Pomphorhynchus laevis reduces lead levels in fish bile, thereby diminishing or impeding the hepatic intestinal cycling of lead, which may reduce the quantity of metals available for fish. Alterations in pollutant uptake and accumulation in different intermediate and final hosts due to parasites are thus very important in the field of ecotoxicology. In addition to such alterations, there is a close interaction between the effects of pollutants and parasites which seems to be mediated at least partly by the endocrine system, which itself is closely related to the immune system in fish. Laboratory studies on eels experimentally infected with the swimbladder nematode Anguillicola crassus reveal that toxic chemicals such as polychlorinated biphenyls produce immunosuppressive effects which facilitate parasite infection. Similarly, an increase in serum cortisol concentration in eels due to chemical exposure and infection is correlated with decreasing levels of anti-A. crassus antibodies. Furthermore, parasites are able to elicit physiological changes which are attributed to chemicals with endocrine disrupting activity, e.g. the cestode Ligula intestinalis is known to suppress gonad development in roach. The most thoroughly documented examples of endocrine disruption in wild fish are in roach, and it is conceivable that this disruption is not only due to chemical activity but also to parasites such as L. intestinalis or species of the phylum Microspora.     

中杜鹃在3种宿主巢中寄生繁殖

了解杜鹃对其宿主的选择和寄生情况,能为两者间的协同进化研究提供重要的基础资料。2012和2013年每年的4～8月,在贵州宽阔水国家级自然保护区对不同生境类型中的鸟巢进行搜索监测,记录到4例中杜鹃(Cuculus saturatus)寄生繁殖现象,其宿主分别是暗绿绣眼鸟(Zosterops japonicus)、棕腹柳莺(Phylloscopus subaffinis)和黄喉鹀(Emberiza elegans),其中棕腹柳莺和暗绿绣眼鸟是首次记录到被中杜鹃寄生。中杜鹃卵重(2.39±0.14)g(n=3),体积(2.24±0.18)cm3(n=3),通过T检验方法发现中杜鹃卵极显著大于暗绿绣眼鸟和棕腹柳莺卵(P0.001),同时在形状上也与两者极不相似,中杜鹃卵呈明显的长椭圆形,与黄喉鹀的卵在大小上无显著差异(P=0.1)。反射光谱的分析结果发现,寄生于不同宿主巢的中杜鹃卵在背景色和斑点上都具有一定差异,这表明中杜鹃的卵可能存在基因族群的分化。     

Modeling the consequence of increased host tolerance toward avian brood parasitism

Avian brood parasites greatly reduce the reproductive success of their hosts. Empirical studies have demonstrated that some hosts have evolved defenses against parasitism like an ability to recognize and reject parasitic eggs that are dissimilar to their own eggs. Detailed mechanisms of how hosts recognize parasitism still remain unknown, but recent studies have shown that the host’s recognition, in many cases, is based on discordance of the eggs in a clutch, and that hosts are more error-prone when the nest is multiply parasitized, i.e., hosts tend to accept more multiple parasitism than single parasitism. In an area in Hungary, the great reed warbler Acrocephalus arundinaceus, one of the main hosts of the common cuckoo Cuculus canorus, is heavily parasitized and the parasitism rate has been kept at quite a high level for decades. Previous mathematical models suggest that such a high parasitism rate can be maintained because the focal host population behaves as a sink where few hosts can reproduce but immigration from outside replenishes the loss of host reproduction in the sink population. Here, we explore the consequences of the increased host tolerance towards multiple parasitism which has been overlooked in the previous studies using a simple model. Our model analysis shows that the increased host tolerance can dramatically contribute to both the parasite abundance and the parasitism rate being kept at a high level. We suggest that such a host behavior, combined with host immigration, can be an important factor responsible for the observed severe parasitism.     

Effects of host age on parasitism and progeny allocation in Trichogrammatidae

The impact of host age on the number of hosts killed, survival of progeny, progeny allocation, and sex allocation was examined for several Trichogrammatidae (Hymenoptera) species in laboratory choice tests. Individual female parasitoids were provided with young, medium-aged and old eggs of one of three lepidopterous host species: Trichoplusia ni (Hübner) (Noctuidae), Pieris rapae (L.) (Pieridae), or Plutella xylostella (L.) (Plutellidae). Trichogrammatid species behaved as gregarious parasitoids with the first two host species, and as solitary parasitoids with eggs of the smaller latter one. They mostly preferred young eggs of T. ni, but did not discriminate among P. rapae eggs of different ages, and often preferred young or medium-aged P. xylostella eggs over old eggs. Survival of progeny did not vary constantly with host age, although it was often very low in P. rapae eggs of any age. Clutch size frequently decreased with host age in both T. ni and P. rapae. Offspring sex ratio did not change with age of T. ni and P. rapae eggs, and rarely did so in P. xylostella eggs. In regard to host age, the results with T. ni are the ones which are the most in agreement with optimal foraging theoretical predictions, as clutch size was the highest in preferred younger eggs.