Characterization of Acromyrmex subterraneus brunneus (Hymenoptera: Formicidae) young nests in a fragment of the Neotropical Forest

Young nests of Acromyrmex subterraneus brunneus are characterized by refuse soil in the exterior of the nest, a single fungus chamber 11 to 20 cm deep in relation to soil surface and internal volume ranging from 0.3 to 1.5 liters. These nidification patterns are important characteristics for identifying and understanding the interactions between species and their habitats.

Internal ambience of bee colonies submitted to strengthening management by adding broods

This study aimed to investigate the ideal brood area to be introduced for strengthening Apis mellifera colonies, considering variations in the hive's internal ambience. Therefore, eight colonies were equipped with data loggers for recording variations in temperature and humidity inside the hives. Then, these colonies were split in four treatments involving the swapping of four, three, two and one brood comb between strong and weak colonies. Growth in the amount of brood and food stores to the four colony strengthening treatments was assessed counting the comb cells to record changes in the brood and food area. The colonies tended to restore their normal thermoregulation within four hours after they had been manipulated and the hive cover had been closed, which quickly increased brood area in the receiving colonies. Results showed that adding up to three brood combs constitute an important alternative for strengthening Apis mellifera colonies because it did not interfere with nest thermoregulation, speeded up population growth in weak colonies and did not affect the development of strong colonies.

Saab J35 Draken -hävittäjän vaikutus Suomen ilmapuolustuskykyyn 1970-luvulla

Kylmän sodan aikana Suomen ilmapuolustus kyseenalaistettiin muiden valtioiden toimesta. Kehittyvän aseteknologian aikana Suomen ilmavoimien hävittäjäkalusto oli jäänyt jälkeen sen ajan sotakalustosta. Tutkattomat hävittäjät kykenivät toteuttamaan ilmavoimille asetettuja tehtäviä vain hyvällä säällä, näkölento-olosuhteissa. Koko valtakunnan ilmapuolustusjärjestelmä koki suuren muutoksen 1970-luvulla, jonka merkittävänä tekijänä oli ruotsalaisen Saab J35 Draken -hävittäjän hankinta ilmavoimien torjuntahävittäjäksi. Tässä tutkimuksessa tarkastellaan uuden torjuntahävittäjän vaikutusta hävittäjätorjunnan toteutukseen ja ilmapuolustuskyvyn ylläpitoon. Tutkimusmenetelmä on laadullinen eli kvalitatiivinen tutkimusmenetelmä, jolloin tutkimuksen aineisto koostuu valtaosin julkaistuista kirjallisuuslähteistä ja haastattelusta. Kaikki suoranaisesti hävittäjätorjuntaa käsittelevät lähteet ovat Draken-aikakaudella ilmavoimissa palvelleiden henkilöiden kirjoittamia. Kirjallisuuslähteiden tueksi haastateltavana henkilönä on käytetty entistä ilmavoimien komentajaa, kenraaliluutnantti Heikki Nikusta. Hankintaprosessin poliittinen valmistelu kesti useita vuosia ja hankintaa jouduttiin lykkäämään eri syistä. Viimein keväällä, vuonna 1970 vahvistettiin kahdentoista Draken-hävittäjän hankinta. Koneet toimitettiin osina Suomeen Valmetin tehtaille koottavaksi. Koneiden oston yhteydessä kauppaan kuului myös koulutuspaketti, jonka avulla tyyppi- ja ilmataistelukoulutus kyettiin aloittamaan Suomessa nopeasti kuuden Ruotsista vuokratun koneen kanssa. Koulutuspaketti edesauttoi Drakenin nopeaa täysimittaista operatiiviseen käyttöön ottoa ja nopeaa ilmataistelukoulutuksen aloitusta Suomessa. Vaikka uusi torjuntahävittäjä oli massiivinen ja hieman kankeaksi kuvailtu sotakone, palautti se hävittäjätorjuntaan sotien jälkeen koulutuksesta kadonneen klassisen ilmataistelun eli kaartotaistelun. Draken oli ensimmäinen hävittäjä Suomen ilmavoimilla, jossa oli tutka ja tutkaohjukset. Oman tutkan käyttöön perustuva ohjushyökkäys paransi hävittäjätorjunnan toteutusta ja samalla se oli täysin uusi koulutettava asia hävittäjäohjaajille. Suuret hävittäjähankinnan kaltaiset sota-asehankinnat ovat merkittäviä ulko- ja turvallisuuspoliittisia päätöksiä. Kylmän sodan aikana Suomen tavoitteena oli edistää puolueetonta ulko- ja turvallisuuspolitiikka. Nootti-kriisi ja läheinen suhde Neuvostoliittoon ei vakuuttanut länsivaltioita Suomen puolueettomuudesta, joten ruotsalaisvalmisteisen torjuntahävittäjän hankinta oli tarkasti harkittu evaluointiprosessissa mukana olleista hävittäjistä. Puolueettomuusimagon lisäksi Draken-hankinta toi mukanaan uusia elementtejä ilmataisteluun. Tutkan ja kaartotaistelun lisäksi uutena mainittavana asiana oli hävittäjäparin käyttö torjuvana yksikkönä. Koneiden pie3 nestä määrästä johtuen Drakenin aikakaudella otettiin käyttöön myös hajautustaktiikka. Koneet sijoitettiin pienempiin osastoihin valtakunnan alueelle, jolloin saatiin kattava tunnistus- ja torjuntakyky ympäri valtakuntaa. Drakenin myötä Suomelle tuli ensimmäistä kertaa historiansa aikana ympärivuorokautinen kyky valvoa ja turvata valtakunnan ilmatilaa sääolosuhteista riippumatta.

Meluntorjunta voiteluainetehtaassa

Melu on epämiellyttävää ja tarpeetonta ääntä. Se saattaa olla myös fysiologisesti haitallista aiheuttaen pahimmassa tapauksessa pysyvän kuulovaurion. Tämän diplomityön tavoitteena oli saada vähennettyä työntekijöiden päivittäistä meluannosta alle 80 desibeliin, jotta työntekijöiden työturvallisuus ja työskentelyolosuhteet paranisivat. Työn teoriaosuudessa kerrotaan yleisiä asioita äänestä, sen mittaamisesta, vaikutuksista ihmisiin, laitevalmistajien ja työnantajien velvollisuuksia sekä meluntorjunnan perusteita. Käytännön osiossa käydään läpi aikaisemmin tehtyjen melumittausten tuloksia, esitellään mittauksissa käytettyjä laitteita, esitellään kohteet joihin parannuksia tehtiin ja miten ne tehtiin, sekä käydään läpi kuinka suuri vaikutus kullakin parannuksella on ollut. Lisäksi verrataan ennen parannuksia tehtyjen meluannosmittausten tuloksia parannusten jälkeisiin tuloksiin. Tulokset olivat hyviä, sillä täyttökoneilla työskentelevien työntekijöiden meluannosmäärät putosivat alle 80 desibeliin.

S500 Suurlujuusteräksestä valmistettujen rakenneputkiliitosten lujuusominaisuudet

Nykyinen Eurocode 3 suunnitteluohjeen rakenneputkiliitoksia käsittelevä osio vaatii S500 suurlujuusteräksestä valmistettujen liitosten mitoituksen yhteydessä käytettäväksi reduk-tiokerrointa 0.8. Tämä on varsin konservatiivinen mitoituspa, joka vähentää merkittävästi lujemman teräslaadun käyttämisestä saavutettavaa hyötyä. Tämän työn pääasiallinen ta-voite on laboratoriotestein selvittää S500 lujuusluokan rakenneputkiliitosten todellinen äärikapasiteetti ja verrata sitä nykyisen mitoitusohjeen mukaiseen kapasiteettiin. Tässä työssä tutkitut liitokset ovat valmistettu poikkileikkaukseltaan nelikulmaisista kyl-mämuovatuista rakenneputkista. Koesarja koostui kahdeksasta X-liitoksesta, sekä kymme-nestä K-liitoksesta. Kaikki testit suoritettiin huoneenlämmössä, LUT Koneen teräsrakenne-laboratoriossa. Laboratoriotestien lisäksi rakenteen käyttäytymistä tutkittiin epälineaarisen elementtimenetelmän avulla ja näitä tuloksia verrattiin kokeellisiin tuloksiin. Sekä testien että FE–analyysin perusteella voidaan todeta, että S500 lujuusluokan terästä käytettäessä ei Eurocode 3:n mukaisen reduktiokertoimen käytölle ole perusteita.

Of milquetoasts and daredevils : personalities in female eiders

Traditionally biologists have often considered individual differences in behaviour or physiology as a nuisance when investigating a population of individuals. These differences have mostly been dismissed as measurement errors or as non-adaptive variation around an adaptive population mean. Recent research, however, challenges this view. While long acknowledged in human personality studies, the importance of individual variation has recently entered into ecological and evolutionary studies in the form of animal personality. The concept of animal personality focuses on consistent differences within and between individuals in behavioural and physiological traits across time and contexts and its ecological and evolutionary consequences. Nevertheless, a satisfactory explanation for the existence of personality is still lacking. Although there is a growing number of explanatory theoretical models, there is still a lack of empirical studies on wild populations showing how traditional life-history tradeoffs can explain the maintenance of variation in personality traits. In this thesis, I first investigate the validity of variation in allostatic load or baseline corticosterone (CORT) concentrations as a measure for differences in individual quality. The association between CORT and quality has recently been summarised under the “CORT-fitness hypothesis”, which states that a general negative relationship between baseline CORT and fitness exists. I then continue to apply the concept of animal personality to depict how the life-history trade-off between survival and fecundity is mediated in incubating female eiders (Somateria mollissima), thereby maintaining variation in behaviour and physiology. To this end, I investigated breeding female eiders from a wild population that breeds in the archipelago around Tvärminne Zoological Station, SW Finland. The field data used was collected from 2008 to 2012. The overall aim of the thesis was to show how differences in personality and stress responsiveness are linked to a life-history context. In the four chapters I examine how the life-history trade-off between survival and fecundity could be resolved depending on consistent individual differences in escape behaviour, stress physiology, individual quality and nest-site selection. First, I corroborated the validity of the “CORT-fitness hypothesis”, by showing that reproductive success is generally negatively correlated with serum and faecal baseline CORT levels. The association between individual quality and baseline CORT is, however, context dependent. Poor body condition was associated with elevated serum baseline CORT only in older breeders, while a larger reproductive investment (clutch mass) was associated with elevated serum baseline CORT among females breeding late in the season. Interestingly, good body condition was associated with elevated faecal baseline CORT levels in late breeders. High faecal baseline CORT levels were positively related to high baseline body temperature, and breeders in poor condition showed an elevated baseline body temperature, but only on open islands. The relationship between stress physiology and individual quality is modulated by breeding experience and breeding phenology. Consequently, the context dependency highlights that this relationship has to be interpreted cautiously. Additionally, I verified if stress responsiveness is related to risk-taking behaviour. Females who took fewer risks (longer flight initiation distance) showed a stronger stress response (measured as an increase in CORT concentration after capture and handling of the bird). However, this association was modulated by breeding experience and body condition, with young breeders and those in poor body condition showing the strongest relationship between risktaking and stress responsiveness. Shy females (longer flight initiation distance) also incubated their clutch for a shorter time. Additionally, I demonstrated that stress responsiveness and predation risk interact with maternal investment and reproductive success. Under high risk of predation, females that incubated a larger clutch showed a stronger stress response. Surprisingly, these females also exhibited higher reproductive success than females with a weaker stress response. Again, these context dependent results suggest that the relationship between stress responsiveness and risk-taking behaviour should not be studied in isolation from individual quality and that stress responsiveness may show adaptive plasticity when individuals are exposed to different predation regimes. Finally, female risk-taking behaviour and stress coping styles were also related to nest-site choice. Less stress responsive females more frequently occupied nests with greater coverage that were farther away from the shoreline. Females nesting in nests with medium cover and farther from the shoreline had higher reproductive success. These results suggest that different personality types are distributed non-randomly in space. In this thesis I was able to demonstrate that personalities and stress coping strategies are persistent individual characteristics, which express measurable effects on fitness. This suggests that those traits are exposed to natural selection and thereby can evolve. Furthermore, individual variation in personality and stress coping strategy is linked to the alternative ways in which animals resolve essential life-history trade-offs.

Biomedical Event Extraction with Machine Learning

Biomedical natural language processing (BioNLP) is a subfield of natural language processing, an area of computational linguistics concerned with developing programs that work with natural language: written texts and speech. Biomedical relation extraction concerns the detection of semantic relations such as protein-protein interactions (PPI) from scientific texts. The aim is to enhance information retrieval by detecting relations between concepts, not just individual concepts as with a keyword search. In recent years, events have been proposed as a more detailed alternative for simple pairwise PPI relations. Events provide a systematic, structural representation for annotating the content of natural language texts. Events are characterized by annotated trigger words, directed and typed arguments and the ability to nest other events. For example, the sentence “Protein A causes protein B to bind protein C” can be annotated with the nested event structure CAUSE(A, BIND(B, C)). Converted to such formal representations, the information of natural language texts can be used by computational applications. Biomedical event annotations were introduced by the BioInfer and GENIA corpora, and event extraction was popularized by the BioNLP'09 Shared Task on Event Extraction. In this thesis we present a method for automated event extraction, implemented as the Turku Event Extraction System (TEES). A unified graph format is defined for representing event annotations and the problem of extracting complex event structures is decomposed into a number of independent classification tasks. These classification tasks are solved using SVM and RLS classifiers, utilizing rich feature representations built from full dependency parsing. Building on earlier work on pairwise relation extraction and using a generalized graph representation, the resulting TEES system is capable of detecting binary relations as well as complex event structures. We show that this event extraction system has good performance, reaching the first place in the BioNLP'09 Shared Task on Event Extraction. Subsequently, TEES has achieved several first ranks in the BioNLP'11 and BioNLP'13 Shared Tasks, as well as shown competitive performance in the binary relation Drug-Drug Interaction Extraction 2011 and 2013 shared tasks. The Turku Event Extraction System is published as a freely available open-source project, documenting the research in detail as well as making the method available for practical applications. In particular, in this thesis we describe the application of the event extraction method to PubMed-scale text mining, showing how the developed approach not only shows good performance, but is generalizable and applicable to large-scale real-world text mining projects. Finally, we discuss related literature, summarize the contributions of the work and present some thoughts on future directions for biomedical event extraction. This thesis includes and builds on six original research publications. The first of these introduces the analysis of dependency parses that leads to development of TEES. The entries in the three BioNLP Shared Tasks, as well as in the DDIExtraction 2011 task are covered in four publications, and the sixth one demonstrates the application of the system to PubMed-scale text mining.

Pup age and aggressive behavior in lactating rats

High levels of aggressive behaviors against intruders in the nest area are displayed by female rats during the first 10 days after delivery, declining thereafter to very low levels, even though lactation continues. Cross-fostering experiments were undertaken to test the hypothesis that pup age may affect aggression in lactating rats. The behavior of females on the 8th day after delivery when raising fostered 8-day-old pups was compared to that of females on the 8th postpartum day raising older pups (18 days old) for the last 5 days, and females on the 18th day after delivery raising fostered 18-day-old pups were compared to females in the same postpartum period nursing younger pups (8 days of age at the time of the maternal aggression test) for 5 days. Pup retrieval activity and plasma prolactin level were also analyzed. Females on the 8th postpartum day nursing 18-day-old pups were less aggressive than females in the same postpartum period, but with 8-day-old pups. Likewise, females on the 18th postpartum day nursing younger pups were more aggressive and presented higher levels of prolactin than females nursing older pups. Thus, pup development can alter the natural decline of maternal aggressive behavior.

Influence of the test situation on pup retrieval behavior of normal and undernourished lactating rats

Undernutrition of dams and pups disrupts the retrieval efficiency of mothers. However, if the mothers are assessed in their home cages, they spend more time with their litters. In the present study the effect of test conditions on pup retrieval behavior of mothers receiving a 25% (well-nourished group) and 8% casein diet (undernourished group) was examined. In agreement with previous studies, undernourished mothers spent more time with their litters than well-nourished dams as lactation proceeded. Pup retrieval behavior varied with test conditions. In the first experiment, the maternal behavior of dams was assessed by the standard procedure (pups were separated from their mother and scattered over the floor of the home cage). The mother was then returned and the number of retrieved pups was recorded. From day 3 to 8, the retrieval efficiency of undernourished dams decreased, while the retrieval efficiency of well-nourished mothers did not vary. In the second experiment, mothers were subjected to a single retrieval test (on day 9 of lactation) using the procedure described for experiment 1. No difference between well-nourished and undernourished mothers was observed. In the third experiment, seven-day-old pups were separated from the mothers and returned individually to a clean home cage. Dietary treatment did not affect the retrieval efficiency. However, undernourished dams reconstructed the nest more slowly than did well-nourished dams. Taken together, these results suggest that pup retrieval behavior of the undernourished mother is not impaired by dietary restriction when the maternal environment is disturbed minimally.

Osuustoiminta ja poliittisuus 1920-luvun Pienviljelijä-lehdessä

Tämän sivuainetutkielman aiheena on pienviljelijäliike. Kysymykseni on, miten pienviljelijäliikettä tuotiin esille Pienviljelijä-lehdessä? Aihetta lähestytään osuustoiminnan ja poliittisuuden kautta. Osuustoiminnan ja poliittisuuden tutkiminen keskittyy tutkimuksessani lehden kyseisistä aiheista käsittelemiin artikkeleihin. Osuustoiminta nähdään tässä tutkimuksessa taloudellisena yhteistoimintana, jossa päämääränä on parantaa siihen osallistuvien taloudellista asemaa. Poliittisuus puolestaan nähdään työväenaatteena, vasemmistolaisuutena. Pienviljelijä on tutkimuksessani itsenäinen maataviljelevä henkilö, joka on torpparivapautuksessa lunastanut muutaman hehtaarin suuruisen maatilan itsensä ja perheensä elatukseen. Olen käyttänyt metodina historiallis-kvalitatiivista eli laadullista tutkimusta pyrkien pelkistämään havaintoja ja sitä kautta ratkaisemaan arvoituksen eli vastaamaan niihin kysymyksiin, joita olen tutkielmalleni asettanut. Tutkimukseni pääasiallinen lähdeaineisto on vuosina 1920—1929 ilmestyneet Pienviljelijä-lehden numerot. Alkuperäisaineiston lisäksi käytän tutkimuskirjallisuutta, josta tärkeimmäksi nousee Arvo Santosen väitöskirja Järjestäytymiskysymys ja pienviljelijäjärjestöjen vakiintuminen Suomessa. Tutkimus maatalouden pienviljelyspoliittisesta murrosvaiheesta 1930-luvun alkuun mennessä. Muuta tutkimuskirjallisuutta olen käyttänyt soveltuvin osin Poliittisuus liittyy tutkimuksessani lehden artikkeleihin, jotka käsittelivät sekä osuus-toimintaa että kunnallis- ja eduskuntavaaleja. Yhteistä molemmissa oli poliittisuuden muuttuminen vuosikymmenen aikana. Vuosikymmenen alussa lehden kannat poliittisuuteen olivat selkeitä, mutta tultaessa vuosikymmenen lopulle niitä pyrittiin häivyttämään ja tilalle tulivat kannanotot, joissa korostettiin pienviljelijäin ammatillisen ja taloudellisen aseman parantamisen merkitystä. Osuustoiminnassa poliittisuus näkyi vaihtoehtona kapitalistiselle talousjärjestelmälle. Osuustoiminnan avulla oli mahdollista saavuttaa sosialistinen järjestelmä. Vaalien alla kirjoittajat olivat huolissaan äänestämisestä ja siitä, että lehden lukijat osasivat valita oikeat ehdokkaat. Alku vuosikymmenellä vaihtoehto ilmaistiin selkeästi, työväen ehdokkaat. Loppuvuosikymmenellä lehdessä ei haluttu selkeästi määrittää ketä tuli äänestää. Lehti ilmoittikin olevansa puolueeton vaalitaisteluiden melskeessä.

Environmental color affects Nile tilapia reproduction

We investigated the effects of environmental color on the reproductive behavior of Nile tilapia, Oreochromis niloticus. Two environmental colors were tested by covering the aquarium (60 x 60 x 40 cm) with white (12 groups) or blue (13 groups) cellophane and observing reproductive behavior in groups of 2 males (10.27 ± 0.45 cm) and 3 females (10.78 ± 0.45 cm) each. After assignment to the respective environmental color (similar luminosity = 100 to 120 Lux), the animals were observed until reproduction (identified by eggs in the female's mouth) or up to 10 days after the first nest building. Photoperiod was from 6:00 h to 18:00 h every day. Food was offered in excess once a day and water quality was similar among aquaria. Daily observations were made at 8:00, 11:00, 14:00 and 17:00 h regarding: a) latency to the first nest, b) number of nests, c) gravel weight removed (the male excavates the nest in the bottom of the aquarium), d) nest area, and e) mouthbrooding incubation (indication of reproduction). The proportion of reproducing fish was significantly higher (6 of 13) in the group exposed to the blue color compared the group exposed to the white color (1 of 12; Goodman's test of proportions). Moreover, males under blue light removed significantly larger masses of gravel (blue = 310.70 ± 343.50 g > white = 130.38 ± 102.70 g; P = 0.01) and constructed wider nests (blue = 207.93 ± 207.80 cm² > white = 97.68 ± 70.64 cm²; P = 0.03) than the control (white). The other parameters did not differ significantly between light conditions. We concluded that reproduction in the presence of blue light was more frequent and intense than in the presence of white light.

Bumblebees in a Changing Climate: Evaluating the Effects of Temperature on Queen Performance

Bumblebees are a very essential group of pollinating insects, but their populations have declined drastically during the past decades. We need to understand why their numbers are decreasing and what can be done to reverse this trend. Climate change-related phenomena, such as changes in the overwintering temperatures and spring conditions, are among the most prominent threats to bumblebees. Queens have a special role in the lifecycle of bumblebees because they overwinter and start new colonies the next year. Their successful performance: survival, overwintering ability, longevity, immune competence, and nest establishing capability in spring, is highly important for bumblebee populations. However, the effects of climate change on bumblebee queen performance remain unknown. The main objective of this thesis was to assess how temperature affects the performance of bumblebee queens during and after overwintering. The effects of warm temperature predicted by climate change scenarios on queen survival and stress-tolerance were studied by a four-month artificial diapause of bumblebee queens at two temperatures (9°C and 1.8°C). Bumblebee colonies were also reared in a laboratory and factors affecting colony characteristics were examined. In addition, queen performance during spring was studied in a starvation experiment using two temperatures (15°C as normal; 24°C as warmer than average) and queens collected from nature right after their emergence. My research revealed how temperature affects queen performance, and queen size was found to be an important factor determining the direction of some of these effects. We found a 0.4g weight threshold for bumblebee queens to be able to survive overwintering. In addition, during mild winters, larger queens have a higher chance than smaller ones to survive through winter and also to cope with immunological stresses after overwintering. During cold conditions, which are normal in the current climatic situation, this advantage disappears. In the spring starvation experiment, the starved queens survived approximately eight days longer in 15°C than in 24°C, which means that starvation risk rises significantly with increasing spring temperature, in a situation where food is scarce due to for example frost damage or asynchrony between bumblebees and their important food plants. These results could mean that in the future climate, larger queens are better able to survive the winter, initiate their nests and start rearing their offspring. This may be problematic, because I also detected two alternative strategies of colony development that differ between large and small queens; larger queens start to lay eggs earlier at nest initiation, their colonies mature later, they produce more workers, and they have a more strongly male biased sex allocation compared with smaller queens. If larger queens have a greater change of producing offspring after a mild winter, this could lead to a significant decline in the total production of new queens at a population level. Thus, it seems that queen size could act as one mechanism regulating the population level outcomes in different temperatures. The new information presented in my thesis reinforces that basic research, monitoring, and local species conservation of bumblebees both in Finland and globally must be increased to ensure that this highly important pollinator group survives in the face of climate change.

Morphine infusions into the rostrolateral periaqueductal gray affect maternal behaviors

It is well established that morphine inhibits maternal behaviors. Previous studies by our group have shown activation of the rostrolateral periaqueductal gray (rlPAG) upon inhibition-intended subcutaneous injections of morphine. In this context, we demonstrated that a single naloxone infusion into the rlPAG, following this opioid-induced inhibition, reactivated maternal behaviors. Since these data were obtained by using peripheral morphine injections, the present study was designed to test whether morphine injected directly into the rlPAG would affect maternal behaviors. Our hypothesis that morphine acting through the rlPAG would disrupt maternal behaviors was confirmed with a local infusion of morphine. The mothers showed shorter latency for locomotor behavior to explore the home cage (P = 0.049). Inhibition was especially evident regarding retrieving (P = 0.002), nest building (P = 0.05) and full maternal behavior (P = 0.023). These results support the view that opioidergic transmission plays a behaviorally meaningful inhibitory role in the rostrolateral PAG.

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