Flut und Hitze: Auswirkungen extremer Klimaereignisse auf die epigäische Arthropodenfauna (Araneae – Spinnen) ufernaher Lebensräume (Auen, Polder) des Inselrheins bei Mainz

Die vorliegende Arbeit ist ein Teil des Projektes Flut und Hitze des Kompetenzzentrums Überflutung an der Universität Mainz. Die Ziele dieser Untersuchung waren: Die Artzusammensetzung und Phänologien der Spinnengemeinschaften von Uferhabitaten bei Mainz (Rheinland-Pfalz, Deutschland) zu ermitteln, anhand des Artenspektrums die Folgen langjähriger Trockenheit und die Auswirkungen des Extremsommers 2003 zu beschreiben, Einflüsse von Überflutungen festzustellen und die Submersionstoleranzen ausgewählter Arten zu bestimmen. Insgesamt wurden 27783 Spinnen aus 179 Arten und 24 Familien bearbeitet. Die Untersuchung umfasste einen Hartholzauwald bei Ingelheim am Rhein, den Hochwasserschutzpolder Ingelheim, sowie Tiermaterial von vier weiteren Uferstandorten und drei Inselstandorten des Rheins bei Mainz. Die Beprobung der Hartholzaue mit Barberfallen und Stammeklektoren erfolgte von Mai 2005 bis Mai 2008. Im Polder wurden von Oktober 2006 bis Mai 2008 mit Barberfallen und einem Vakuumsauger gefangen. Die Proben der weiteren Standorte stammten aus Barberfallenfängen der Jahre 2000 bis einschließlich 2005. In der seit Winter 2002/2003 nicht mehr überfluteten und im Sommer stark austrocknenden Hartholzaue wurde eine als xerotolerant zu bezeichnende Spinnenfauna vorgefunden. Dies galt insbesondere für die sehr artenreiche Stammregion. Zu den dominierenden Spezies zählten: Diplostyla concolor (Boden), Clubiona pallidula und Textrix denticulata (beide Stamm). Der Polder Ingelheim wurde überwiegend von euryöken Freilandbewohnern besiedelt, dominant kamen Oedothorax apicatus und Pardosa agrestis vor. Das Tiermaterial der Ufer- und Inselstandorte wies deutliche Unterschiede in der Artenzusammensetzung im Bezug auf die Flutungsintensität auf. Nach dem Ausbleiben von Hochwässern und dem starken Austrocknen der Standorte im Sommer 2003 wurden hygrobionte Arten wie Allomengea vidua kaum mehr vorgefunden, während sich xerotolerante Spezies ausbreiteten. Darüber hinaus wurden die Submersionstoleranzen ausgewählter Spinnenarten im Labor ermittelt. Die gewonnenen Daten lassen Vermuten, dass die getesteten Spinnenspezies durchaus in der Lage sind, bei niedrigen Wassertemperaturen eine kurze Flut submers zu überstehen. Unter hohen Temperaturen besteht dagegen keine Submersionstoleranz.

Comparative analysis of midline guidance and axon pathfinding in the chelicerates Achaearanea tepidariorum and Cupiennius salei

Die Neurogenese und axonale Wegfindung sind in den vergangenen Jahrzehnten Thema einer Vielzahl wissenschaftlicher Untersuchungen in den verschiedensten Organismen gewesen. Die zusammengetragenen Daten in Insekten und Crustaceen geben eine gute Übersicht darüber, wie das Nervensystem in Arthropoden aufgebaut wird. Die entwicklungsbiologischen Prozesse, die daran beteiligt sind, sind in den beiden genannten Gruppen sehr gut verstanden. In den Gruppen der Cheliceraten und Myriapoden jedoch wurden ähnliche Analysen bisher kaum durchgeführt. Das Hauptanliegen dieser Arbeit war es daher, Mechanismen in den Spinnen Achaearanea tepidariorum und Cupiennius salei, zwei Vertretern der Cheliceraten, zu untersuchen, die eine Rolle im Leitsystem der ventralen Mittellinie und bei der axonalen Wegfindung spielen. Eine Vorraussetzung hierfür sind Kenntnisse über die Architektur des Zentralnervensystems. In einem ersten Schritt beschrieb ich daher grundlegend die Morphologie des Nervensystems im Verlauf der gesamten Embryoalentwicklung. Ich konnte zeigen, dass in Spinnen ein für Arthropoden typisches Strickleiternervensystem gebildet wird. Dieses wird von segmental angelegten Neuronen geformt, wobei sowohl Gruppen von Zellen als auch einzelne Neurone daran beteiligt sind, die primären axonalen Trakte zu etablieren. Im Besonderen konnte ich eine Zelle identifizieren, die in Position, Projektionsmuster und der Expression des Markergens even-skipped vergleichbar zum PR2 Neuron in Drosophila ist, welches die posteriore Wurzel des Segmentalnervs anlegt.rnrnIn einem zweiten Ansatz untersuchte ich die ventrale Mittellinie in Spinnen im Bezug auf ihre mögliche Funktion in der axonalen Wegfindung. Es konnte gezeigt werden, dass es sich beim Epithel der Mittellinie, das die Lücke zwischen beiden Keimstreifhälften während des gesamten Prozesses der Inversion überspannt, um eine transiente Struktur handelt, die keine neuralen Zellen hervorbringt. Es ist daher vergleichbar mit der so genannten Floor plate in Vertebraten, die ebenfalls nur vorübergehend existiert. Die Untersuchung von single minded (sim) zeigte, dass es, anders als in Drosophila, wo sim ein wichtiges regulatorisches Gen für die korrekte Spezifizierung von Mittellinienzellen ist, nicht in den Zellen der Mittellinie, sondern in diesen benachbarten Zellen, exprimiert wird. Das ist vergleichbar mit Vertebraten. Zusätzlich konnte ich Expression von sim an den Basen der Gliedmassen und im Kopf nachweisen. Wie in Vertebraten könnte sim an der Musterbildung dieser Gewebe beteiligt sein. Dennoch spielt die Mittellinie in Spinnen eine wichtige Rolle als Organisator für auswachsende, kommissurale Axone. Diese Funktion teilt sie mit anderen Invertebraten und Vertebraten.rnrnDie Signaltransduktionskaskade, die an der axonalen Wegfindung an der Mittellinie beteiligt ist, ist in den verschiedensten Organismen hoch konserviert. In der vorliegenden Arbeit konnte ich sowohl in Achaearanea als auch in Cupiennius ein netrin Homolog identifizieren und eine konservierte Funktion des Wegfindungsmoleküls während der Bildung der Kommissuren aufzeigen. RNAi Experimente belegen, dass, wird die Funktion von netrin herunterreguliert, das Strickleiternervensystem nicht korrekt gebildet wird, ins Besondere die kommissuralen Faszikel. Des Weiteren konnte ich eine neue Funktion von netrin, die bisher in anderen Organsimen noch nicht beschrieben wurde, identifizieren. Neben seiner Rolle in der axonalen Wegfindung, scheint netrin auch an der epithelialen Morphogenese im zentralen Nervensystem beteiligt zu sein. In dieser Funktion scheint netrin in Gliazellen, die die epithelialen Vesikel der Invaginationsgruppen umhüllen, wichtig zu sein, um neurale Vorläuferzellen in einem undifferenzierten Zustand zu halten. Der Abbau von netrin Transkript durch RNA Interferenz führt zu einer verfrühten Segregation neuraler Vorläuferzellen aus dem epithelialen Verband der Invaginationsgruppen und zu einer Zunahme an Zellen, die den frühen Differenzierungsmarker islet exprimieren.

Funktionelle Charakterisierung der Metalloendopeptidasen Meprin alpha und Meprin beta unter besonderer Berücksichtigung pathologischer Umstände in der menschlichen Dermis

Im Rahmen dieser Arbeit wurden zwei neuentdeckte Astacin-ähnliche-Proteasen LAST undrnLAST_MAM aus dem Pfeilschwanzkrebs Limulus polyphemus funktionell charakterisiert.rnInsbesondere LAST_MAM, eignet sich zur phylogenetischen Untersuchung, hinsichtlich derrnEvolution von Astacin-ähnlichen-Proteasen mit MAM-Domäne, zu denen auch die Meprinernzählen. Es wurde deutlich, dass LAST_MAM nicht unmittelbar mit anderen Astacinen, diernüber eine MAM-Domäne verfügen, verwandt ist, und dass von einer divergenten Entwicklungrndieser Proteasen und der MAM-Domäne selbst ausgegangen werden muss.rnMeprin Metalloendopeptidasen werden in membrangebundener und sezernierter Form,rnvorwiegend in Epithelien aber auch in intestinalen Leukozyten und bestimmten Krebszellenrnexprimiert. Meprin

Performance studies of the CMS distributed analysis system in the associated Higgs boson production with top quarks

L'obiettivo di questa tesi è studiare la fattibilità dello studio della produzione associata ttH del bosone di Higgs con due quark top nell'esperimento CMS, e valutare le funzionalità e le caratteristiche della prossima generazione di toolkit per l'analisi distribuita a CMS (CRAB versione 3) per effettuare tale analisi. Nel settore della fisica del quark top, la produzione ttH è particolarmente interessante, soprattutto perchè rappresenta l'unica opportunità di studiare direttamente il vertice t-H senza dover fare assunzioni riguardanti possibili contributi dalla fisica oltre il Modello Standard. La preparazione per questa analisi è cruciale in questo momento, prima dell'inizio del Run-2 dell'LHC nel 2015. Per essere preparati a tale studio, le implicazioni tecniche di effettuare un'analisi completa in un ambito di calcolo distribuito come la Grid non dovrebbero essere sottovalutate. Per questo motivo, vengono presentati e discussi un'analisi dello stesso strumento CRAB3 (disponibile adesso in versione di pre-produzione) e un confronto diretto di prestazioni con CRAB2. Saranno raccolti e documentati inoltre suggerimenti e consigli per un team di analisi che sarà eventualmente coinvolto in questo studio. Nel Capitolo 1 è introdotta la fisica delle alte energie a LHC nell'esperimento CMS. Il Capitolo 2 discute il modello di calcolo di CMS e il sistema di analisi distribuita della Grid. Nel Capitolo 3 viene brevemente presentata la fisica del quark top e del bosone di Higgs. Il Capitolo 4 è dedicato alla preparazione dell'analisi dal punto di vista degli strumenti della Grid (CRAB3 vs CRAB2). Nel capitolo 5 è presentato e discusso uno studio di fattibilità per un'analisi del canale ttH in termini di efficienza di selezione.

Megistobenthic faunal diversity of the Antalya Gulf: Crustacea

The spatio-temporal distribution of megistobenthic crustacean assemblages from the Antalya Gulf, located in the Levantine Sea is described. In order to provide a comprehensive overview of the spatio-temporal patterns of the crustacean community, 3 transect including depth of 10, 25, 75, 125 and 200 m, were studied between 2014 and 2015 to investigate their association with a set of environmental parameters in representative months of each season (spring, summer, autumn and winter). For its economic importance in Levantine waters, a focus analysis of deep-water rose shrimp Parapenaeus longirostris (Lucas, 1846) was done, to investigate the length frequency composition of the population of the Antalya Gulf. A total of 58 crustacean species were encountered in the study area, of these species identified, 18 species were recognized as alien species in the Mediterranean Sea. Throughout the year the most frequent species of the study were the hermit crab Pagurus prideaux (Leach, 1815) and Parapenaeus longirostris (Lucas, 1846) followed by the Indo-Pacific swimming crab Charybdis longicollis (Leene, 1938) and by the invasive shrimp Marsupenaeus japonicus (Spence Bate, 1888). Few species contributing to a high amount to the total biomass were found throughout the year. These species were Charybdis longicollis and Parapenaeus longirostris. Stations of the study area showed similar values of diversity indices of benthic crustacean community among the three transect. The highest values of faunistic indices were detected in autumn and winter (October and February), and also varied along the depth gradient, with the highest values found between 25 and 75 meters. The multivariate analyses conducted on the abundance data point out major differences between depths and between seasons. Therefore, according to cluster analysis and ordination over abundance and biomass, three main crustacean assemblages were detected: the first corresponding to shallow bottoms (10, 25 meters), the second corresponding to intermediate waters (75 meters) and the last to deeper waters (125, 200 meters). Depth was the main factor governing the distribution of megistobenthic crustacean in the area. Besides the depth, the structure of the sediment is the most important factor in determining the crustacean assemblage. Therefore, all factors governing the crustacean distribution were found to be related to the bottom depth. The population of Parapenaeus longirostris in the Antalya Gulf showed significant differences in depth. It was found that females dominated the population of the study area (65.11%), and were significantly larger than males for each cohort identified. The size-weight relationships revealed a slight negative allometry in growth, a bit more pronounced in females than in males.

Purification, cDNA structure and biological significance of a single insulin-like growth factor-binding domain protein (SIBD-1) identified in the hemocytes of the spider Cupiennius salei

Cupiennius salei single insulin-like growth factor-binding domain protein (SIBD-1), which exhibits an IGFBP N-terminal domain-like profile, was identified in the hemocytes of the spider C. salei. SIBD-1 was purified by RP-HPLC and the sequence determined by a combination of Edman degradation and 5'-3'- RACE PCR. The peptide (8676.08 Da) is composed of 78 amino acids, contains six intrachain disulphide bridges and carries a modified Thr residue at position 2. SIBD-1 mRNA expression was detected by quantitative real-time PCR mainly in hemocytes, but also in the subesophageal nerve mass and muscle. After infection, the SIBD-1 content in the hemocytes decreases and, simultaneously, the temporal SIBD-1 expression seems to be down-regulated. Two further peptides, SIBD-2 and IGFBP-rP1, also exhibiting IGFBP N-terminal domain variants with unknown functions, were identified on cDNA level in spider hemocytes and venom glands. We conclude that SIBD-1 may play an important role in the immune system of spiders.

Species Richness-Environment Relationships of European Arthropods at Two Spatial Grains: Habitats and Countries

We study how species richness of arthropods relates to theories concerning net primary productivity, ambient energy, water-energy dynamics and spatial environmental heterogeneity. We use two datasets of arthropod richness with similar spatial extents (Scandinavia to Mediterranean), but contrasting spatial grain (local habitat and country). Samples of ground-dwelling spiders, beetles, bugs and ants were collected from 32 paired habitats at 16 locations across Europe. Species richness of these taxonomic groups was also determined for 25 European countries based on the Fauna Europaea database. We tested effects of net primary productivity (NPP), annual mean temperature (T), annual rainfall (R) and potential evapotranspiration of the coldest month (PETmin) on species richness and turnover. Spatial environmental heterogeneity within countries was considered by including the ranges of NPP, T, R and PETmin. At the local habitat grain, relationships between species richness and environmental variables differed strongly between taxa and trophic groups. However, species turnover across locations was strongly correlated with differences in T. At the country grain, species richness was significantly correlated with environmental variables from all four theories. In particular, species richness within countries increased strongly with spatial heterogeneity in T. The importance of spatial heterogeneity in T for both species turnover across locations and for species richness within countries suggests that the temperature niche is an important determinant of arthropod diversity. We suggest that, unless climatic heterogeneity is constant across sampling units, coarse-grained studies should always account for environmental heterogeneity as a predictor of arthropod species richness, just as studies with variable area of sampling units routinely consider area.

To look or not to look at threat? Scanpath differences within a group of spider phobics

Predicting the behavior of phobic patients in a confrontational situation is challenging. While avoidance as a major clinical component of phobias suggests that patients orient away from threat, findings based on cognitive paradigms indicate an attentional bias towards threat. Here we present eye movement data from 21 spider phobics and 21 control subjects, based on 3 basic oculomotor tasks and a visual exploration task that included close-up views of spiders. Relative to the control group, patients showed accelerated reflexive saccades in one of the basic oculomotor tasks, while the fear-relevant exploration task evoked a general slowing in their scanning behavior and pronounced oculomotor avoidance. However, this avoidance strongly varied within the patient group and was not associated with the scores from spider avoidance-sensitive questionnaire scales. We suggest that variation of oculomotor avoidance between phobics reflects different strategies of how they cope with threat in confrontational situations.

Hypervigilance-avoidance pattern in spider phobia

Cognitive-motivational theories of phobias propose that patients' behavior is characterized by a hypervigilance-avoidance pattern. This implies that phobics initially direct their attention towards fear-relevant stimuli, followed by avoidance that is thought to prevent objective evaluation and habituation. However, previous experiments with highly anxious individuals confirmed initial hypervigilance and yet failed to show subsequent avoidance. In the present study, we administered a visual task in spider phobics and controls, requiring participants to search for spiders. Analyzing eye movements during visual exploration allowed the examination of spatial as well as temporal aspects of phobic behavior. Confirming the hypervigilance-avoidance hypothesis as a whole, our results showed that, relative to controls, phobics detected spiders faster, fixated closer to spiders during the initial search phase and fixated further from spiders subsequently.

A two year study of verified spider bites in Switzerland and a review of the European spider bite literature

During a two-year study, all spider bites recorded by Swiss primary care physicians were reported to the Swiss Toxicological Information Centre and all collected spiders were identified. A total of 14 verified spider bites were recorded, involving five species from four families: Zoropsis spinimana (five cases), Cheiracanthium punctorium (four cases), Tegenaria atrica (three cases) and one case of Malthonica ferruginea (¼ Tegenaria ferruginea) (both Agelenidae), and one case of Amaurobius ferox (Amaurobiidae). The bites of all spider species produced relatively mild symptoms. Local symptoms such as moderate to severe pain, circumscribed swelling and redness were the only effects in most cases. Systemic symptoms were rare. There was complete recovery in all cases and all lesions healed completely without further damage or secondary disorders. Following a review of the European spider bite literature, the number of spider species capable of biting humans in Europe is considered to be much larger than could be concluded from this study. Most spider bites are restricted to species living synanthropically, thus promoted by climate and habitat change. The annual frequency of spider bites in Switzerland is estimated at 10 – 100 bites per million inhabitants, but this is predicted to increase due to the continuous arrival of new alien species, many of which have a high potential to establish in urban areas

Effects of Habitat Fragmentation on Abundance, Larval Food and Parasitism of a Spider-Hunting Wasp.

Habitat fragmentation strongly affects species distribution and abundance. However, mechanisms underlying fragmentation effects often remain unresolved. Potential mechanisms are (1) reduced dispersal of a species or (2) altered species interactions in fragmented landscapes. We studied if abundance of the spider-hunting and cavity-nesting wasp Trypoxylon figulus Linnaeus (Hymenoptera: Crabronidae) is affected by fragmentation, and then tested for any effect of larval food (bottom up regulation) and parasitism (top down regulation). Trap nests of T. figulus were studied in 30 agricultural landscapes of the Swiss Plateau. The sites varied in the level of isolation from forest (adjacent, in the open landscape but connected, isolated) and in the amount of woody habitat (from 4 % to 74 %). We recorded wasp abundance (number of occupied reed tubes), determined parasitism of brood cells and analysed the diversity and abundance of spiders that were deposited as larval food. Abundances of T. figulus were negatively related to forest cover in the landscape. In addition, T. figulus abundances were highest at forest edges, reduced by 33.1% in connected sites and by 79.4% in isolated sites. The mean number of spiders per brood cell was lowest in isolated sites. Nevertheless, structural equation modelling revealed that this did not directly determine wasp abundance. Parasitism was neither related to the amount of woody habitat nor to isolation and did not change with host density. Therefore, our study showed that the abundance of T. figulus cannot be fully explained by the studied trophic interactions. Further factors, such as dispersal and habitat preference, seem to play a role in the population dynamics of this widespread secondary carnivore in agricultural landscapes.

Spider Ecophysiology

With over 43,000 species, spiders are the largest predacious arthropod group. They have developed key characteristics such as multi-purpose silk types, venoms consisting of hundreds of components, locomotion driven by muscles and hydraulic pressure, a highly evolved key-lock mechanism between the complex genital structures, and many more unique features. After 300 million years of evolutionary refinement, spiders are present in all land habitats and represent one of the most successful groups of terrestrial organisms. Ecophysiology combines functional and evolutionary aspects of morphology, physiology, biochemistry and molecular biology with ecology. Cutting-edge science in spiders focuses on the circulatory and respiratory system, locomotion and dispersal abilities, the immune system, endosymbionts and pathogens, chemical communication, gland secretions, venom components, silk structure, structure and perception of colours as well as nutritional requirements. Spiders are valuable indicator species in agroecosystems and for conservation biology. Modern transfer and application technologies research spiders and their products with respect to their value for biomimetics, material sciences, and the agrochemical and pharmaceutical industries.

Spider Venoms Potentially Lethal to Humans

Spiders have one pair of venom glands, and only a few families have reduced them completely (Uloboridae, Holarchaeidae) or modified them to another function (Symphytognathidae or Scytodidae, see Suter and Stratton 2013). All other 42,000 known spider species (99%) utilize their venom to inject it into prey items, which subsequently become paralysed or are killed. Spider venom is a complex mixture of hundreds of components, many of them interacting with cell membranes or receptors located mainly in the nervous or muscular system (Herzig and King 2013). Spider venom, as it is today, has a 300-million-yearlong history of evolution and adaptation and can be considered as an optimized tool to subdue prey. In Mesothelae, the oldest spider group with less than 100 species, the venom glands lie in the anterior part of the cheliceral basal segment. They are very small and do not support the predation process very effectively. In Mygalomorphae, the venom glands are well developed and fill the basal cheliceral segment more or less completely. Many of these 3,000 species are medium- to large-/very large-sized spiders, and they have created the image of being dangerous beasts, attacking and killing a variety of animals, including humans. Although this picture is completely wrong, it is persistent and contributes considerably to human arachnophobia. The third group of spiders, Araneomorphae or “modern spiders”, comprises 93% of all spider species. The venom glands are enlarged and extend to the prosoma; the openings of the venom ducts are moved from the convex to the concave side of the cheliceral fangs and enlarged as well. These changes save the chelicerae from the necessity of being large, and hence, on the average, araneomorph spiders are much smaller than mygalomorphs. Nevertheless, they possess relatively large venom glands, situated mainly in the prosoma, and may also have rather potent venom.

Main Components of Spider Venoms

Venom glands are alreadypresent in theoldes t spider group, the Mesothelae. Theglands lie in the anterior portion of the cheliceral basal segment but are very small, and it is doubtful how much the venom contributes to the predatory success. In mygalomorph spiders, the well-developed venom glands are still in the basal segment of the chelicerae and produce powerful venom that is injected via the cheliceral fangs into a victim. In all other spiders (Araneomorphae), the venom glands have become much larger and reach into the prosoma where they can take up a considerable proportion of this body part. Only a few spiders have reduced their venom glands, either partially or completely (Uloboridae, Holarchaeidae and Symphytognathidae are usually mentioned) or modified them significantly (Scytodidae, see Suter and Stratton 2013). As well as using venom, spiders may also use their chelicerae to overwhelm an item of prey. It is primarily a question of size whether a spider chews up small arthropods without applying venom or if it injects venom first. Very small and/or defenceless arthropods are picked up and crashed with the chelicerae, while larger, dangerous or well-defended items are carefully approached and only attacked with venom injection. Some spiders specialize on prey groups, such as noctuid moths (several genera of bola spiders among Araneidae), web spiders (Mimetidae), ants (Zodarion species in Zodariidae, aphantochiline thomisids, several genera among Theridiidae, Salticidae, Clubionidae and Gnaphosidae) or termites (Ammoxenidae). However, these more or less monophagous species amount only to roughly 2 % of all known spider species, while 98 % are polyphagous. From these considerations, it follows that the majority of spider venoms are not tailored to any given invertebrate or insect group but are rather unspecialized to be effective over a broad spectrum of prey types that spiders naturally encounter.

Glucocorticoids enhance in vivo exposure-based therapy outcome in spider phobia

BACKGROUND: Preclinical and clinical studies indicate that the administration of glucocorticoids may promote fear extinction processes. In particular, it has been shown that glucocorticoids enhance virtual reality based exposure therapy of fear of heights. Here, we investigate whether glucocorticoids enhance the outcome of in vivo exposure-based group therapy of spider phobia. METHODS: In a double blind, block-randomized, placebo-controlled, between-subject study design, 22 patients with specific phobia of spiders were treated with two sessions of in vivo exposure-based group therapy. Cortisol (20 mg) or placebo was orally administered 1 hr before each therapy session. Patients returned for a follow-up assessment one month after therapy. RESULTS: Exposure-based group therapy led to a significant decrease in phobic symptoms as assessed with the Fear of Spiders Questionnaire (FSQ) from pretreatment to immediate posttreatment and to follow-up. The administration of cortisol to exposure therapy resulted in increased salivary cortisol concentrations and a significantly greater reduction in fear of spiders (FSQ) as compared to placebo at follow-up, but not immediately posttreatment. Furthermore, cortisol-treated patients reported significantly less anxiety during standardized exposure to living spiders at follow-up than placebo-treated subjects. Notably, groups did not differ in phobia-unrelated state-anxiety before and after the exposure sessions and at follow-up. CONCLUSIONS: These findings indicate that adding cortisol to in vivo exposure-based group therapy of spider phobia enhances treatment outcome.