Can MRI predict olfactory loss and improvement in posttraumatic olfactory dysfunction?

BACKGROUND: Although most patients with post-traumatic olfactory dysfunction (PTOD) undergo MRI, there is no consensus about its diagnostic or prognostic value. The aims were: 1) to classify the extent of post-traumatic neurodegeneration; 2) to determine its relationship with chemosensory dysfunction (smell, taste, trigeminal); and 3) to establish whether MRI can predict olfactory improvement. METHODOLOGY: We conducted a retrospective cohort study based on a series of 56 patients with PTOD. All patients underwent validated psychophysical tests of their smell, taste, and trigeminal functions, otorhinolaryngologic evaluation, and MRI. An experienced radiologist blinded to patient data evaluated 40 chemosensory-relevant brain regions according to a four-point scale (0=no lesion to 3=large lesion). Follow up data after 4 years (on average) were available in 46 patients. RESULTS: The cluster analysis showed 4 brain lesion patterns that differed in lesion localization and severity. They are associated with diagnostic categories: anosmia, hyposmia and normosmia. Two clusters were highly specific for anosmia (100% specificity)and could accurately predict this condition (100% positive predictive value). No clusters were associated with trigeminal or taste dysfunction. Regarding improvement, 72.7% of patients in the cluster with mild lesions experienced subjective and measurable olfactory improvement whereas this was only the case in 21.7-37.5% of patients with larger lesions. The odds of subjective smell improvement were 5.9 times higher in patients within the milder cluster compared to larger ones. CONCLUSIONS: The analysis of brain lesions in PTOD allows corroboration of smell test results and prediction of subjective and measurable improvement.

Effects of COVID-19 on the Human Central Olfactory System: A Natural Pre-Post Experiment.

BACKGROUND AND PURPOSE: Reduced olfactory function is the symptom with the highest prevalence in coronavirus disease 2019 (COVID-19) with nearly 70% of infected individuals experiencing partial or total loss of their sense of smell at some point during the disease. The exact cause is not known, but beyond peripheral damage, studies have demonstrated insults to both the olfactory bulb and central olfactory brain areas. However, these studies often lack both baseline pre-COVID-19 assessments and control groups, and the effects could, therefore, simply reflect pre-existing risk factors. MATERIALS AND METHODS: Shortly before the COVID-19 outbreak, we completed an olfactory-focused study, which included structural MR brain images and a full clinical olfactory test. Opportunistically, we invited participants back 1 year later, including 9 participants who had experienced mild-to-moderate COVID-19 (C19+) and 12 who had not (C19-), creating a natural pre-post experiment with a control group. RESULTS: Despite C19+ participants reporting subjective olfactory dysfunction, few showed signs of objectively altered function. Critically, all except 1 individual in the C19+ group had reduced olfactory bulb volume (average reduction, 14.3%), but this did not amount to a significant statistical difference compared with the control group (2.3%) using inference statistics. We found no morphologic differences in olfactory brain areas but stronger functional connectivity between olfactory brain areas in the C19+ group at the postmeasure. CONCLUSIONS: Our data suggest that COVID-19 might cause long-term reduction in olfactory bulb volume and altered functional connectivity but with no discernible morphologic differences in cerebral olfactory regions.

Hedonic perception of odors in children aged 5-8 years is similar across 18 countries: Preliminary data.

OBJECTIVE: Olfactory preference emerges very early in life, and the sense of smell in children rapidly develops until the second decade of life. It is still unclear whether hedonic perception of odors is shared in children inhabiting different regions of the globe. METHODS: Five-hundred ten healthy children (N = 510; ngirls = 256; nboys = 254) aged from 5 to 8 years from 18 countries rated the pleasantness of 17 odors. RESULTS: The hedonic perception of odors in children aged between 5 and 8 years was rather consistent across 18 countries and mainly driven by the qualities of an odor and the overall ability of children to label odorants. CONCLUSION: Conclusions from this study, being a secondary analysis, are limited to the presented set of odors that were initially selected for the development of U-Sniff test and present null findings for the cross-cultural variability in hedonic perception of odors across 18 countries. These two major issues should be addressed in the future to either contradict or replicate the results presented herewith. This research lays fundament for posing further research questions about the developmental aspects of hedonic perception of odors and opens a new door for investigating cross-cultural differences in chemosensory perception of children.

Central olfactory processing in patients with disorders of consciousness.

BACKGROUND AND PURPOSE: Previous studies have demonstrated that individuals suffering from disorder of consciousness (DOC) maintain some minor neural processing of percepts mediated by senses that early in their pathway intersect the thalamus, a key dysfunctional area in DOC patients. Here the degree of sensory preservation within the olfactory system, a system that lacks an obligatory thalamic relay, and its relationship to the consciousness level in DOC patients of various etiologies was assessed. METHODS: Clinical Coma Recovery Scale - Revised (CRS-R) as well as cerebral responses to odors by means of functional magnetic resonance were obtained in a group of vegetative state/unresponsive wakefulness syndrome (n = 26) patients, minimally conscious state (n = 7) patients and healthy controls (n = 25). RESULTS: A majority of vegetative state/unresponsive wakefulness syndrome patients (58%) and 100% of minimally conscious state patients demonstrated a significant preservation of olfactory neural processing, manifested by activation within the piriform cortex, an area considered as a primary olfactory region. Degree of preservation of olfactory processing differed linearly in line with the patients' etiologies where groups demonstrating greater conscious awareness demonstrated more significant processing. Viewed over all DOC patients, there was a significant negative association between odor-related activity in the orbitofrontal cortex and CRS-R scores. CONCLUSIONS: It is demonstrated that DOC patients exhibit a significant preservation of olfactory neural processing with a clear relationship to etiopathologies and clinical measures even years after of chronification of DOC.

Insect emergence in relation to floods in wet meadows and swamps in the River Dalälven floodplain.

Annual variation in flood frequency and hydroperiod during the vegetation season has ecological impacts on the floodplain biota. Although many insect groups may have a lower emergence during a flood event, it is poorly known how annual emergence of insects in temporary wetlands is related to the variation in hydrology. Between May and September, we studied the weekly emergence of 18 insect taxa over six consecutive years, 2002-2007, in six temporary flooded wetlands (four wet meadows and two forest swamps) in the River Dalälven floodplains, Central Sweden. We used emergence traps to collect emerging insects from terrestrial and aquatic parts of wet meadows and swamp forests. In all wetlands, the insect fauna was numerically dominated by the orders Diptera, Hymenoptera, Coleoptera and Homoptera. On a weekly basis, 9 out of the 18 insect taxa had lower emergence in weeks with flood than in weeks with no flood, whereas no taxon had a higher emergence in weeks with flood. Over the seasons, we related insect emergence to seasonal flood frequency and length of hydroperiod. The emergence of most studied taxa decreased with increasing hydroperiod, which suggests that emergence after floods do not compensate for the reduced emergence during floods. Only Culicidae and the aquatic Chironomidae sub-families Tanypodinae and Chironominae showed an increase in emergence with increasing hydroperiod, whereas Staphylinidae peaked at intermediate hydroperiod. We conclude that a hydroperiod covering up to 40% of the vegetation season has a significant negative effect on the emergence of most taxa and that only a few taxa occurring in the temporary wetlands are actually favoured by a flood regime with recurrent and unpredictable floods.

Francisella tularensis subspecies holarctica occurs in Swedish mosquitoes, persists through the developmental stages of laboratory-infected mosquitoes and is transmissible during blood feeding.

In Sweden, mosquitoes are considered the major vectors of the bacterium Francisella tularensis subsp. holarctica, which causes tularaemia. The aim of this study was to investigate whether mosquitoes acquire the bacterium as aquatic larvae and transmit the disease as adults. Mosquitoes sampled in a Swedish area where tularaemia is endemic (Örebro) were positive for the presence of F. tularensis deoxyribonucleic acid throughout the summer. Presence of the clinically relevant F. tularensis subsp. holarctica was confirmed in 11 out of the 14 mosquito species sampled. Experiments performed using laboratory-reared Aedes aegypti confirmed that F. tularensis subsp. holarctica was transstadially maintained from orally infected larvae to adult mosquitoes and that 25% of the adults exposed as larvae were positive for the presence of F. tularensis-specific sequences for at least 2 weeks. In addition, we found that F. tularensis subsp. holarctica was transmitted to 58% of the adult mosquitoes feeding on diseased mice. In a small-scale in vivo transmission experiment with F. tularensis subsp. holarctica-positive adult mosquitoes and susceptible mice, none of the animals developed tularaemia. However, we confirmed that there was transmission of the bacterium to blood vials by mosquitoes that had been exposed to the bacterium in the larval stage. Taken together, these results provide evidence that mosquitoes play a role in disease transmission in part of Sweden where tularaemia recurs.

The arbovirus vector Culex torrentium is more prevalent than Culex pipiens in northern and central Europe.

Two species of arbovirus vector, Culex torrentium and Culex pipiens (Diptera: Culicidae), occur in several European countries, but difficulties in their accurate identification and discrimination have hampered both detailed and large-scale distribution and abundance studies. Using a molecular identification method, we identified to species 2559 larvae of Cx. pipiens/torrentium collected from 138 sites in 13 European countries ranging from Scandinavia to the Mediterranean coast. In addition, samples of 1712 males of Cx. pipiens/torrentium collected at several sites in the Czech Republic were identified to species based on the morphology of their hypopygia. We found that the two species occur together in large areas of Europe, and that Cx. torrentium dominates in northern Europe and Cx. pipiens dominates south of the Alps. The transition in dominance occurs in central Europe, where both species are roughly equally common. There was a strong correlation between the length of the growing season at different sites and occurrences of the two species. As the growing season increases, the proportion and detection of Cx. torrentium decrease, whereas those of Cx. pipiens increase. The present findings have important consequences for the interpretation of the results of studies on major enzootic and link-vectors of mosquito-borne bird-associated viruses (i.e. Sindbis, West Nile and Usutu viruses), especially in central Europe and Scandinavia.

Gustatory and olfactory dysfunction in older adults: a national probability study.

BACKGROUND: Olfactory and gustatory functions have not been well characterized in older adults in the US. Consequently, their relationships to sociodemographic characteristics, as well as physical and mental health, were studied in a large national probability sample using brief validated tests of chemosensory function. METHODS: A five-odour identification test and taste-impregnated strips of filter paper (sweet, sour, bitter, and salty) assessed the ability to identify chemosensory stimuli. RESULTS: Severe gustatory dysfunction was more prevalent than severe olfactory dysfunction. Age, education and sex were independently associated with performance on both the olfactory and gustatory identification tasks. Higher scores were associated with female sex, higher level of education, and lower age. Odour identification scores exhibited a positive, albeit weak, correlation with BMI, and food-related odours were better identified than non-food odours. In addition, odour identification performance was also negatively associated with depressive symptoms. CONCLUSIONS: These data demonstrate a high prevalence of severe gustatory and, to a somewhat lesser extent, olfactory dysfunction in a population-based sample and demonstrate that even brief tests are capable of detecting correlations between both chemical senses and relevant health measures outside a clinical setting.

Perception of specific trigeminal chemosensory agonists.

The intranasal trigeminal system is a third chemical sense in addition to olfaction and gustation. As opposed to smell and taste, we still lack knowledge on the relationship between receptor binding and perception for the trigeminal system. We therefore investigated the sensitivity of the intranasal trigeminal system towards agonists of the trigeminal receptors TRPM8 and TRPA1 by assessing subjects' ability to identify which nostril has been stimulated in a monorhinal stimulation design. We summed the number of correct identifications resulting in a lateralization score. Stimuli were menthol (activating TRPM8 receptors), eucalyptol (TRPM8), mustard oil (TRPA1) and two mixtures thereof (menthol/eucalyptol and menthol/mustard oil). In addition, we examined the relationship between intensity and lateralization scores and investigated whether intensity evaluation and lateralization scores of the mixtures show additive effects. All stimuli were correctly lateralized significantly above chance. Across subjects the lateralization scores for single compounds activating the same receptor showed a stronger correlation than stimuli activating different receptors. Although single compounds were isointense, the mixture of menthol and eucalyptol (activating only TRPM8) was perceived as weaker and was lateralized less accurately than the mixture of menthol and mustard oil (activating both TRPM8 and TRPA1) suggesting suppression effects in the former mixture. In conclusion, sensitivity of different subpopulations of trigeminal sensory neurons seems to be related, but only to a certain degree. The large coherence in sensitivity between various intranasal trigeminal stimuli suggests that measuring sensitivity to one single trigeminal chemical stimulus may be sufficient to generally assess the trigeminal system's chemosensitivity. Further, for stimuli activating the same receptor a mixture suppression effect appears to occur similar to that observed in the other chemosensory systems.

A sensitive and reliable restriction enzyme assay to distinguish between the mosquitoes Culex torrentium and Culex pipiens.

Culex pipiens pipiens Linnaeus and Culex torrentium Martini (Diptera: Culicidae) are closely related vector species that exist sympatrically in Europe. The two species are morphologically almost identical and can only be distinguished with certainty by characters of the male genitalia. Hence, correct species identification and conclusions on distribution and vector status are very difficult and often neglected. Therefore, we developed a reliable and simple mitochondrial cytochrome c oxidase subunit I (COI) gene restriction enzyme assay to discriminate between Cx. pipiens and Cx. torrentium, based on the analysis of morphologically identified male specimens. We sequenced approximately 830 bp in the 3' region of the mitochondrial COI gene of 18 morphologically identified males of Cx. pipiens and Cx. torrentium. Two restriction enzymes (FspBI and SspI) that could distinguish between the two species according to species-specific differences in these sequences were chosen. The restriction enzymes were tested on 227 samples from Sweden and verified by sequencing 44 of them. The enzyme FspBI correctly identified all investigated samples; the enzyme SspI identified all samples except one Cx. torrentium. We hope the method and the findings presented here will help to shed light on the true distribution and relative proportions of the two species in Europe.

A six-year study of insect emergence from temporary flooded wetlands in central Sweden, with and without Bti-based mosquito control.

In temporary wetlands in the River Dalälven floodplains, recurrent but irregular floods induce massive hatching of the flood-water mosquito Aedes sticticus, which causes enormous nuisance. Flood-water mosquito control using the biological larvicide Bacillus thuringiensis var. israelensis (Bti) was commenced in parts of the floodplains during 2002, and here we report the first six years of full-season monitoring of general insect emergence from temporary wetlands with and without treatment. Emergence traps, which were emptied weekly, were used from May to September each year. A total of 137,153 insects of 13 taxonomic orders were collected. Diptera was highly dominating and especially the sub-order Nematocera with 18 families was a very prominent taxon. Bti-treatment effects were analysed by taxonomic order, by sub-order in Diptera and Hemiptera, and by family for Nematocera and Coleoptera for the whole study period. We found no significant negative effects of Bti treatments on the production of insects by taxonomic order, with the exception of Coleoptera in the long term. However, no significant negative effects were found for the Coleoptera families, neither in the short term nor in the long term. There was no significant negative treatment effect on Nematocera production, neither when analyzed for the whole sub-order nor when analyzed by family. However, abundance of Ceratopogonidae was significantly higher in experimental than in reference wetlands. We conclude that Bti-treatment effects on insect production may be minute in comparison to other environmental factors structuring the insect fauna of the temporary wetlands studied.

The fish is bad: Negative food odors elicit faster and more accurate reactions than other odors.

Dissociating between 'good' or 'bad' odors is arguable of crucial value for human survival, since unpleasant odors often signal danger. Therefore, negative odors demand a faster response in order to quickly avoid or move away from negative situations. We know from other sensory systems that this effect is most evident for stimuli from ecologically-relevant categories. In the olfactory system the classification of odors into the food or non-food category is of eminent importance. We therefore aimed to explore the link between odor processing speed and accuracy and odor edibility and valence by assessing response time and detection accuracy. We observed that reaction time and detection accuracy are influenced by both pleasantness and edibility. Specifically, we showed that an unpleasant food odor is detected faster and more accurately than odors of other categories. These results suggest that the olfactory system reacts faster and more accurately to ecologically-relevant stimuli that signal a potential danger.

Production of wetland Chironomidae (Diptera) and the effects of using Bacillus thuringiensis israelensis for mosquito control.

Massive mosquito nuisance problems, caused by the flood-water mosquito Aedes sticticus, occur after floods in the flood plains of the River Dalälven, central Sweden. Since 2002, the biological mosquito larvicide Bacillus thuringiensis var. israelensis (Bti) has been used to control these mosquitoes. Here, we report results from the first six years of monitoring Chironomidae, the most susceptible non-target organisms, in three wetlands with Bti-treatment against mosquitoes and in three wetlands without treatment. Emergence traps were used for continuous insect sampling from May to September each year, 2002-2007, and were emptied once a week. A total of 21,394 chironomids of 135 species were collected, and the subfamily Orthocladiinae dominated the fauna followed by Chironominae and Tanypodinae. The annual chironomid production in the wetlands was low, with an average of 1917 individuals per m(2), and 42 g ash-free dry weight per m(2). We found no reduced production of chironomids at neither family, nor subfamily level, in Bti-treated as compared to untreated wetlands. This is the first long-term follow-up study of the possible effects of Bti-based mosquito larval control on chironomid species production. In the short-term view, one species had higher production in treated areas. In the long-term view, four species had higher and one species had lower production in treated areas. We conclude that the Bti-based control of flood-water mosquitoes does not cause any major direct negative effects on chironomid production, and therefore does not seem to induce any risk for indirect negative effects on birds, bats or any other predators feeding on chironomids.

High species richness of Chironomidae (Diptera) in temporary flooded wetlands associated with high species turn-over rates.

Species richness and species turn-over of Chironomidae was studied in irregularly flooded wetlands of the River Dalälven flood-plains in central Sweden. The chironomid fauna, sampled with emergence traps in six wetlands over six summers, contained as much as 135 species, and the cumulative species curves indicated that the regional species pool contain several more species. Recurrent irregular floods may have induced this high chironomid species richness and the high species turn-over in the temporary wetlands, as the dominance between terrestrial and aquatic species shifted between years. Half of the wetlands were treated with Bacillus thuringiensis var. israelensis (Bti) against larvae of the flood-water mosquito Aedes sticticus. These treatments had no significant effect on chironomid species richness, but there was a higher species turn-over between years of primarily low abundance species in the treated wetlands. The cumulative number of species was also higher in the Bti-treated experimental wetlands than in the untreated reference wetlands. Thus, Bti treatment against mosquito larvae seemed to have only small effects on chironomid species richness but seemed to increase the colonisation-extinction dynamics.

Comparison of mosquito (Diptera: Culicidae) populations by wetland type and year in the lower river Dalälven region, Central Sweden.

We studied adult mosquito assemblages in six wetlands, representing three types (wet meadow, alder swamp, and bog), in the lower part of the River Dalälven in Central Sweden during three consecutive years (2000-2002) and evaluated the influence of wetland type and year. Mosquito abundance differed significantly between years but not between wetland types. Mosquito species richness showed no significant variation between years or wetland types. Cluster analysis based on percentage of similarity resulted in three clusters, with high similarity between all wetlands in 2000. Ordination analysis showed that mosquito assemblages were mainly correlated with wetland type and water level increase in the previous month. Hydrological conditions varied between the years and between the wetland types, and our collections also included a year (2000) with extreme flood situations. The floodwater mosquito species Ochlerotatus sticticus was the predominant species with a strong influence on the whole study due to its long-range dispersal ability. The entire region suffered from enormous numbers of Oc. sticticus in 2000. The data from this study provided the basis for the initiation of a mosquito control project in the region.

Different responses of two floodwater mosquito species, Aedes vexans and Ochlerotatus sticticus (Diptera: Culicidae), to larval habitat drying.

Insect larvae that live in temporary ponds must cope with a rapidly diminishing resource. We tested the hypothesis that floodwater mosquitoes would react to diminishing water levels by accelerating larval development time and emerging as smaller adults. Since a reduction in habitat size leads to increased larval densities, we also included two larval densities. Newly-hatched floodwater mosquito larvae, Aedes vexans (87.9% of emerged adults) and Ochlerotatus sticticus (12.0% of emerged adults), were taken from the field and randomly assigned to one of three water level schedules. Survival to adult emergence was significantly affected by the water level schedule. Ae. vexans adults emerged later in the decreasing schedule than the constant water schedule, but time to emergence was not affected by larval density. In the drying water schedule, Ae. vexans adults emerged 6 to 14 days after complete water removal. Adult size was significantly affected by both water level schedule and larval density. Adults of Oc. sticticus emerged earlier in the decreasing than the constant water schedule which was in accordance with our hypothesis, but size was not affected. Our results indicate two different responses of two floodwater mosquito species to diminishing larval habitat. Oc. sticticus accelerated larval development while Ae. vexans larvae showed remarkable survival in humid soil. Both species are often numerous in inundation areas of large rivers, and climatic conditions after a flood might influence which species dominates the adult mosquito fauna.

Biological diversity versus risk for mosquito nuisance and disease transmission in constructed wetlands in southern Sweden.

In southern Sweden, many wetlands have been constructed, and maintaining or increasing biological diversity is often included in the aims. Some wetlands are constructed near human settlements, thus raising the problem of wetlands being associated with mosquitoes (Diptera: Culicidae). Increased biodiversity (including mosquito diversity) is considered desirable, whereas mosquito nuisance from a human point of view is not. Adult mosquito abundance, diversity and species assemblages of constructed wetlands were compared to natural wetlands. The potential of constructed wetlands for mosquito nuisance and transmission of mosquito-borne viruses was evaluated. The study areas included five constructed and four natural wetlands. Mosquito abundance and species richness were higher in the natural than in the constructed wetlands, and showed a positive correlation with wetland size. Mosquito species assemblages formed three clusters, which were not explained by origin, size and water permanence of wetlands. In a redundancy analysis, however, mosquito faunas showed significant relationships with these variables, and size and origin of wetlands were most important. Major nuisance species (multivoltine species feeding on mammals and laying eggs on soil) were found in all wetlands, although in relatively low numbers. Risk assessment for Sindbis virus transmission showed moderate risk for two constructed wetlands near human settlements. It is concluded that small size of constructed wetlands has the advantage of low mosquito numbers from a human point of view. The use of functional groups is recommended as a tool for presenting mosquito data to the public, and for helping communication between scientists and administrative decision makers.

Pcons: a neural-network-based consensus predictor that improves fold recognition.

During recent years many protein fold recognition methods have been developed, based on different algorithms and using various kinds of information. To examine the performance of these methods several evaluation experiments have been conducted. These include blind tests in CASP/CAFASP, large scale benchmarks, and long-term, continuous assessment with newly solved protein structures. These studies confirm the expectation that for different targets different methods produce the best predictions, and the final prediction accuracy could be improved if the available methods were combined in a perfect manner. In this article a neural-network-based consensus predictor, Pcons, is presented that attempts this task. Pcons attempts to select the best model out of those produced by six prediction servers, each using different methods. Pcons translates the confidence scores reported by each server into uniformly scaled values corresponding to the expected accuracy of each model. The translated scores as well as the similarity between models produced by different servers is used in the final selection. According to the analysis based on two unrelated sets of newly solved proteins, Pcons outperforms any single server by generating approximately 8%-10% more correct predictions. Furthermore, the specificity of Pcons is significantly higher than for any individual server. From analyzing different input data to Pcons it can be shown that the improvement is mainly attributable to measurement of the similarity between the different models. Pcons is freely accessible for the academic community through the protein structure-prediction metaserver at http://bioinfo.pl/meta/.

Prevalence of sindbis virus neutralizing antibodies among Swedish passerines indicates that thrushes are the main amplifying hosts.

The Sindbis virus uses birds as vertebrate hosts in the summer amplification cycle, and the virus is transmitted by ornithophilic Culex species. Previous field and experimental studies have shown that mainly passerine birds are involved in the amplification. To delineate the pattern of Sindbis virus infections among passerines, we collected and sampled birds for blood at five study sites located in northern, central, and southern Sweden. All study sites were lowland forested wetlands and humid forests. The blood samples were assayed for Sindbis neutralizing antibodies, and we tested if the prevalence of Sindbis antibodies varied in relation to bird characteristics (i.e., species, body-mass, sex, and age), and environmental factors (i.e., year, month, and location). We found that Sindbis virus infections occurred in almost all passerine species sampled, but that the infection prevalence was unequally distributed among species. The fieldfare, the redwing, and the songthrush each had significantly higher prevalence than the average for all species. Large passerine species had higher infection prevalence than small species. The infection was less prevalent in hatching-year birds than in older birds during June and July, but not in August. Males and females had the same infection prevalence. The prevalence of Sindbis antibodies was higher in central than in southern Sweden, which coincided with a higher proportion of fieldfare-redwing-songthrush samples in the central region of the country. Thus, it is possible that regional and annual variations in the prevalence of Sindbis antibodies in Swedish passerine species depend on the number of fieldfares, redwings, and songthrushes available for feeding by vector mosquitoes.

The effects of testosterone on a viral infection in greenfinches (Carduelis chloris): an experimental test of the immunocompetence-handicap hypothesis.

The immunocompetence-handicap hypothesis suggests that the honesty of quality signals could be guaranteed if testosterone (T) suppresses immune function while enhancing male ornaments. In addition, it has been proposed that the cost of enhancing ornaments should be highest for males with small ornaments. Recently, the assertion that T causes obligate immunosuppression has been questioned. In this study, we tested whether elevated T levels would increase susceptibility to a viral infection, and whether this hypothesized effect would be most pronounced in males with small ornaments. We surgically inserted T implants into 15 male greenfinches (Carduelis chloris) and control implants into a further 15 males. All birds were then infected with a naturally occurring virus (Sindbis virus, Alphavirus genus), and each bird's daily viraemia (blood virus concentration) was measured for seven days. The specific antibody response was measured for eight weeks. T-implanted males did not exhibit increased viraemia or decreased antibody response, and males with small and large ornaments did not respond differently to T implantation. We did, however, find that T implantation decreased viraemia early in the course of the infection and increased viraemia late in the infection. Thus, our results demonstrate that T may act both to increase and to decrease viraemia.