Berlin, Freie University, Diss., 2005. / Dateiformat: zip, Dateien im PDF-Format.

In dieser Arbeit untersuche ich das Verhalten von Arbeiterbienen beim Brutwärmen, die Wärmeübertragung von den Bienen auf die gedeckelte Brut, die thermophysikalischen Eigenschaften des Brutnests und spezielle Aspekte des Brutnestaufbaus, die für dieses Thema relevant sind und bisher nicht untersucht wurden. Meine Arbeit umfasst Verhaltensbeobachtungen und thermografische Messungen an individuellen Bienen, die Simulation des Heizverhaltens von Arbeiterinnen und das Messen der Temperaturänderungen in der Wabe, die Messung der thermophysikalischen Eigenschaften der Brutwabe und der Zellwände (Wärmeleitfähigkeit und Durchlässigkeit für Wärmestrahlung), die Auswertung von Brutzelltemperaturen als Ergebnis des Verhaltens von Arbeiterbienen, die Analyse der Anzahl und der räumlichen Verteilung von Brutlücken (Auswertung in 2-D und 3-D bezüglich beider Wabenseiten) und die Entwicklung spezifischer Computersoftware, die zur Erarbeitung dieser Ergebnisse unverzichtbar ist. Ein wichtiges Ergebnis dieser Arbeit ist die Entdeckung und Beschreibung eines bemerkenswerten, bislang unbekannten Verhaltens der Honigbiene: Die Aufrechterhaltung hoher Thoraxtemperaturen (TTh) bei Langzeitbesuchen in offenen Zellen („Lücken“) die verstreut in der gedeckelten Brutfläche vorkommen. Hier zeige ich, dass die Aufrechterhaltung der hohen TTh nicht auf den Zellinhalt (z. B. offene Brut) bezogen ist - in den meisten Fällen waren die besuchten Zellen ohnehin leer - sondern auf die direkt benachbarte gedeckelte Brut, mit der diese Zellen über gemeinsame Zellwände in Kontakt stehen. Dieses Verhalten liefert eine Erklärung für Langzeitzellbesuche von sehr langer Dauer ohne erkennbare Aktivität, die in früheren Arbeiten beschrieben aber nicht völlig verstanden wurden, und es rehabilitiert die scheinbar „faulen“ Bienen im Zellinnern. Diesem Verhalten kommt eine große Bedeutung für das Brutwärmen zu, da sich der aufgeheizte Thorax tief in der Wabe (fast an der Mittelwand) befindet wo der Wärmeverlust an die Luft minimiert ist und von wo bis zu 6 umliegende Puppenzellen gleichzeitig gewärmt werden können. Im Vergleich zum Brutwärmeverhalten an der Wabenoberfläche (Andrücken des Thorax an die Brutdeckel), wo nur 1 oder Teile von 3 Brutdeckeln mit dem Thorax in Berührung stehen, ist das Wärmen im Zellinnern mit derselben TTh bis zu 2,6-fach effizienter. Die Messung der thermophysikalischen Eigenschaften der Brutwabe und die Simulation des Brutwärmeverhaltens unter kontrollierten Bedingungen zeigen, dass sich die Wabe langsam aufwärmt und eher ein lokal begrenztes Wärmen als eine rasche Wärmeausbreitung über eine große Fläche begünstigt. Der Einflussbereich eines einzelnen Zellbesuchers hängt von seiner TTh und der Dauer des Zellbesuchs ab. Anstiege der Bruttemperatur in bis zu 3 Zellen Abstand zum Zellbesucher sind nachweisbar. Das hier beschriebene Brutwärmeverhalten im Innern von Lücken (offenen Zellen) bietet nicht nur neue Einsichten in das Bienenverhalten. Es ermöglicht auch eine Neubewertung der Lücken und ihrer Nützlichkeit für die Bienen. Eine von mir entwickelte Computersoftware („CombUse 2.0“) ermöglicht es, das Vorkommen und die räumliche Verteilung von Lücken mit hoher Genauigkeit auf der Ebene einzelner Zellen zu erfassen und auszuwerten. Die räumliche Verteilung der Lücken in der gedeckelten Brutfläche zeigt, dass schon bei geringen Lückenhäufigkeiten von ca. 4 bis 10 %, die in gesunden Kolonien normal sind, eine überraschend große Zahl gedeckelter Brutzellen (88 % bis 99 %, wenn die dreidimensionale Verteilung berücksichtigt wird) im Einflussbereich von Brut wärmenden Zellbesuchern sind. Obwohl das Brutwärmeverhalten im Zellinnern schwer zu entdecken und zu beobachten ist, führen die in dieser Arbeit präsentierten Daten zu dem Schluss, dass es sich dabei um einen wichtigen Bestandteil der Nestklimatisierung bei Honigbienen handelt. / In this thesis I investigate the behaviour of worker bees while incubating brood, heat transfer from bees to the sealed brood, the thermophysical properties of the brood nest and special aspects of brood nest architecture that are relevant for this topic but have not been investigated so far. My work comprises behavioural observations and thermographic measurements of individual bees, simulation of worker bees’ heating behaviour and measurement of temperature changes within the brood comb, the measurement of thermophysical properties of the brood comb and cell walls (thermal conductivity and transmission of thermal radiation), the analysis of brood cell temperatures as a result of the behaviour of worker bees, the analysis of the number and spatial distribution of brood gaps (analysis in 2-D and 3-D with regard to both sides of a comb) and the engineering of specific computer software which was indispensable to achieve these results. One major result of this work is the discovery and description of a very remarkable, hitherto unknown behaviour of the honeybee: the maintenance of high thorax temperatures (TTh) during long-time visits to open cells (“gaps”) which are scattered at low rates among the sealed brood. Here I show that the maintenance of high TTh is not related to the contents of the visited cells (anyway, they were empty in most cases) but to the presence of sealed brood which is directly adjacent to and sharing common cell walls with the visited cell. The discovery of this behaviour casts a positive light on apparently “lazy” bees and provides an explanation for long-time cell-visits with durations of several ten minutes which were described but not fully understood in earlier works by other authors. Moreover, this behaviour is of high relevance for brood incubation itself because the heated thorax is deep in the comb (almost down to the middle wall) where heat loss to the air is minimized and where up to 6 surrounding pupa cells can be warmed simultaneously. In comparison to specific brood incubation behaviour on the comb surface (pressing the thorax onto the brood caps), where only one or parts of three brood caps may be in touch with a heated thorax, heating inside cells with the same TTh is up to 2,6 times more efficient. The measurement of thermophysical properties of the brood comb and the simulation of worker heating under controlled conditions show that the comb warms up slowly and supports local heating rather than a rapid spread of heat all over the area. The radius-of-influence of a single cell visitor depends on its TTh and on the duration of the cell visit. Increases of brood temperature as far as three cells away from the cell visitor may be detected. The brood incubation behaviour via gaps (i.e. open cells) which I describe in this work does not only give new insights into bee behaviour. It also allows a reconsideration of the gaps them¬selves and their usefulness to the bees. A specific computer software (“CombUse 2.0”) which I developed for this work allowed me to register and analyze the spatial distribution of gaps with high precision on the level of cells, in contrast to rough estimations of brood areas and gap numbers which are commonly used in bee-breeding and population assessments. The analysis of the spatial distribution of gaps in the sealed brood area shows that even at small proportions of gaps (ca. 4 to 10 %, common to healthy colonies), a surprisingly large number (88 % to 99 %, if the three-dimensional distribution of gaps is considered) of sealed brood cells is within the reach of brood incubating worker bees visiting these gaps. Although brood incubation behaviour inside cells is very difficult to detect and observe, the data presented in this work strongly suggest that heating inside cells is an important part of nest climate control and other aspects of social life in honeybee colonies.

<p>Morfometrijske analize su ra&ntilde;ena merenjem krilne nervature 540 uzoraka krila<br />pčela sa 9 lokaliteta u Vojvodini. Izračunato je 16 uglova (A1, A4, B3, B4, D7, E9, G7,<br />G18, H12, J10, J16, K19, L13, M17, O26, Q21) koje zaklapa krilna nervatura i 4 indeksa<br />(Ci, Pci, Dbi, Ri), ukupno 20 mera. Izračunate su prosečne vrednosti i utvr&ntilde;ena je<br />statistička značajnost razlika izme&ntilde;u pčela iz regona Srema i Bačke i pčela iz regiona<br />Banata, tako&ntilde;e je izvr&scaron;eno i pore&ntilde;enje pčela svih lokaliteta sa DAWINO standardima za<br />5 rasa pčela (Apis mellifera carnica, Apis mellifera macedonica, Apis mellifera mellifera,<br />Apis mellifera ligustica i Apis mellifera caucasica).<br />Analizom varijanse izračunatih 20 osobina krilne nervature, utvr&ntilde;eno je da samo<br />kod osobine A4 nisu utvr&ntilde;ene statistički značajnie razlike izme&ntilde;u posmatranih<br />lokaliteta, dok su u 19 osobina utvr&ntilde;ene statistički značajne razlike.<br />Utvr&ntilde;ivanjem statističke značajnosti razlika, pčela iz regiona Srema i Bačke i pčela<br />iz regiona Banata, utvr&ntilde;eno je da 45% osobina ne pokazuju statistički značajne razlike,<br />dok 45% osobina pokazuje statistički vrlo značajne razlike (P&lt;0,01) i 10% osobina<br />pokazuje statistički značajne razlike (P&lt;0,05).<br />Upore&ntilde;ivanjem dobijenih vrednosti 20 parametera krilne nervature, pomoću ztesta,<br />sa DAWINO standardima za pet rasa pčela, utvr&ntilde;eno je da na bazi celog uzorka<br />statistički nema značajnih razlika kod osobina A4 i D7 sa A. m. carnica, kod osobina<br />H12, G18 i B4 sa A. m.macedonica i kod osobina J16 i B4 pore&ntilde;eno sa rasom A.<br />m.ligustica. Kod pčela iz regiona Srema i Bačke utvr&ntilde;eno je da statistički nema značajnih<br />razlika kod osobina A4, B3, D7 i G18 upore&ntilde;eno sa A.m. carnica, kod osobina H12 i B4<br />upore&ntilde;eno sa A. m.macedonica i kod osobina G18, K19, J16 i Q21 upore&ntilde;eno sa rasom<br />A. m.ligustica, dok kod pčela iz regiona Banata utvr&ntilde;eno je da statistički nema značajnih<br />razlika kod osobina A4, E9, D7 i J10 upore&ntilde;eno sa A.m. carnica, kod osobina H12, J10,<br />L13 i PCi upore&ntilde;eno sa A. m.macedonica i kod osobina B4, J16 i PCi upore&ntilde;eno sa<br />rasom A. m.ligustica.<br />Ocena genetičke povezanosti, unutar populacijska raznolikost i struktura<br />populacije, dva tipa pčela u Vojvodini, izračunata je na bazi varijacije alela 25 lokusa<br />mikrosatelita. Izvr&scaron;ena je genetska tipizacija sledećih mikrosatelita: A8, A14, A24, A29,<br />A43, A79, A88, A113, Ac11, Ac88, Ac139, Ac306, Ap15, Ap68, Ap85, Ap90, Ap223,<br />Ap224, Ap226, Ap249, Ap273, Ap274, Ap288, At168, At188. 92% ili 23 lokusa su se<br />pokazali kao polimorfni u uzorcima pčela iz Srema i Bačke, a 88% ili 22 lokusa su se<br />pokazali kao polimorfni u uzorcima pčela iz Banata. Izračunata heterozigotnost na nivou<br />cele populacije se nije statistički značajno razlikovala od očekivane heterozigotnosti.<br />Utvr&ntilde;eno je da dobijene genetičke razlike izme&ntilde;u analiziranih pčela iz regiona Srema i<br />Bačke i retgiona Banata nisu dovoljne da se ove dve populacije mogu smatrati<br />razdvojenim.</p> / <p> Morphometric analyses have been done by measuring the wing nervature in<br /> 540 samples of bees, collected from nine localities in Vojvodina. 16 angles<br /> formed by wing nervation have been calculated(A1, A4, B3, B4, D7, E9, G7,<br /> G18, H12, J10, J16, K19, L13, M17, O26, Q21) as well as four indexes (Ci, PCI,<br /> DBI , R), a total of 20 measures. The average values have been calculated and<br /> statistical significant differences in bees from Srem, Backa and Banat region<br /> determined. Five breeds of bees from these regions have been compared to<br /> Dawino standards.<br /> The analyses of the variance of calculated 20 features of wing nervature indicate<br /> that statistically significant differences in monitored localities have not been found<br /> only in A4, on the other hand in 19 properties significant differences have been<br /> discovered.<br /> Established statistically significant differences between breeds from Srem<br /> and Backa regions reveale that 45% properties do not show any statistically<br /> important differences, while 45% features show very important statistical<br /> differences (P&lt;0,01) and 10% show statistically important differences (P&lt;0,05).<br /> It has been established by comparing the obtained values of 20 parametres<br /> of wing nervature by means of z test to DAWINO standards for five breeds of<br /> bees that, based on the whole sample, there are no significant differences in<br /> features A4 and D7 in A.m. carnica, in features H12, G18 and B4 in A.m.<br /> macedonica and features J16 and B4 compared to A.m. ligustica. As for bees from<br /> Srem and Backa region,there are statistically no significante differences in<br /> features A4, B3, D7 and G18 compared to A.m. carnica, features H12 and B4<br /> compared to A.m. macedonica and features G18, K19, J16 and Q21 compared to<br /> A.m. ligustica, while in bees from Banat region, statistically there are no<br /> significant differences in features A4, E9, D7 and J10 compared to A.m. carnica,<br /> features H12, J10, L13 and Pci compared to A.m. macedonica and features B4,<br /> J16 and Pci compared to A.m. ligustica.<br /> The evaluation of genetic correlation, the diversity of bees population and<br /> population structure of two types of bees in Vojvodina have been established on<br /> the basis of allels variations in 25 locus microsatelites.The following<br /> microsatelites have been standardized &ndash; A8,A14,A24,A29, A43, A79, A88, A113,<br /> Ac11, Ac88, Ac139, Ac306, Ap15, Ap68, Ap85, Ap90, Ap223, Ap224, Ap226,<br /> Ap249, Ap273, Ap274, Ap288, At168, At188. 92% or 23 locus have shown as<br /> polymorphs in bees from Srem and Backa and 88% or 22 locus samples have<br /> shown as polzmorphs in bees samples from Banat and Backa region. The whole<br /> population calculated heterozygosity has not shown statistically significant<br /> differrence from expected heterozygisity. It has been established that the obtained<br /> genetic differences between the analysed bees from Srem and Backa region and<br /> Banat region are not significant to indicate two populations.</p>

<p>Dokazivanje virusnih infekcija pĉela od velikog je interesa za pravilnu procjenu odnosa izmeĊu domaćina i potencijalnog patogena. Do izrade ove doktorske teze, na teritoriji Bosne i Hercegovine nije bilo prouĉavanja koja bi dokaz virusnih infekcija pĉela dovela do mogućnosti kliniĉke procjene patolo&scaron;kog odnosa za eko-genotip prilagoĊene pĉelinje zajednice. Evolucijski adaptirani virusi medonosne pĉele Apis mellifera carnica, sa stanovi&scaron;ta veterinarske entomologije, veterinarske virusologije i infektivnih bolesti, te saznanja veterinarskog servisa uop&scaron;te, nisu bili dovoljno prepoznati. Pojava i dostupnost molekularnih dijagnostiĉkih metoda omogućile su senzitivno i specifiĉno dokazivanje dijela genoma virusa koji inficiraju pĉelinje zajednice. Iz ekstrahovanih uzoraka templata razliĉitih uzoraka homogenizata sluĉajno uzorkovanih pĉela, RNK, sa jednim parom prajmera za svaki virus, pojedinaĉno, redoslijedom: BQCV, DWV, SBV, ABPV, KBV, CBPV uraĊeni su amplifikacijski protokoli i dobijeni rezultati za klasiĉan, end point, RT-PCR. Ovom doktorskom disertacijom dobijeni su prvi rezultati koji se odnose na utvrĊivanje prisustva i ra&scaron;irenosti virusnih infekcija kod vrste Apis mellifera u pĉelinjacima Bosne i Hercegovine. Od oko dvadeset virusa koji inficiraju pĉele i pĉelinje zajednice, metodama primjenjive molekularne dijagnostike u ovoj doktorskoj tezi dokazivali smo prisustvo virusnih infekcija za &scaron;est najvi&scaron;e prouĉavanih virusa pĉela. Dobijeni rezultati dokazali su prisustvo pet od &scaron;est traţenih virusa sa razliĉitim kombinacijama koinfekcija te raznolikom prostornom distribucijom virusa. Poznavanje sloţenih mehanizama virusnih infekcija pĉelinjih zajednica neophodnost je koja sa novim saznanjima progla&scaron;ava neke od virusa emergentnim patogenima, koji bitno naru&scaron;avaju opstanak pĉelinje zajednice. U patolo&scaron;ko-sinergistiĉkom odnosu sa nametnikom pĉelinje zajednice Varroa destructor neki pĉelinji virusi uzroci su globalno registrovanih gubitaka. Dio istraţivanja u radu odnosio se na terensko prouĉavanje dinamike prisustva grinje Varroa destructor u pĉelinjim zajednicama. Za pravilno dono&scaron;enje suda o uzroku ugibanja pĉelinjih zajednica, pored postojećih zahtjeva kontrole patogena pĉela definisanih procedurama dijagnostiĉkog manuala i Koda OIE-a, bilo bi potrebno imati i rezultate kojim bi se dokazalo kvalitativno i kvantitativno prisustvo virusa kod vrste Apis mellifera. UvoĊenje i standardizacija molekularno-biolo&scaron;kih metoda u rutinskoj dijagnostici virusnih bolesti pĉela je opravdano. Samo je pitanje vremena kada će virusne infekcije pĉela biti predmet rutinske dijagnostike kojim se kontroli&scaron;ezdravstveni status pĉelinje zajednice.</p> / <p>Evidence of viral infection of bees is of great interest for the proper assessment of the relationship between host and pathogen potential. By making this doctoral thesis on the territory of Bosnia and Herzegovina, where before was no present study of evidence of viral infections, lead to ability on clinical evaluation of pathological relations for eco-genotype adapted colonies. Evolutionarily adapted viruses honeybee Apis mellifera, from the veterinary entomology, veterinary virology and infectious diseases, and knowledge of the veterinary services in general, have not been recognized enough. The appearance and availability of molecular diagnostic methods have allowed sensitive and specific detection part of the genome of the virus that infects the colonies. Templates were extracted from samples of different patterns, homogenisate, randomly sampled bees, RNA, with a pair of primers for each virus, respectively, in the order: BQCV, DWV, SBV, ABPV, KBV, CBPV amplification protocols were done and the results obtained by conventional, end point, RT -PCR. This doctoral dissertation obtained the first results concerning the determination of the presence and spread of viral infections in the species Apis mellifera in Bosnia and Herzegovina. From about twenty viruses that can infect bees and hives using diagnostic methods applicable in this doctoral thesis we demonstrate the presence of viral infections of six most studied viruses of bees. The obtained results showed the presence five of the six essential viruses in forms of co-infection with different combinations and varied virus spatial distribution. Awareness of the complex mechanisms of viral infections in colonies is that the new knowledge proclaimed a virus as &quot;emergent pathogen&quot; that significantly impairs the survival of honeybee colonies. The pathogenic synergistic relationship with the parasite Varroa destructor causes that bee viruses are globally listed as reasons for losses. Part of the research work was related to the field study of the dynamics of the presence of Varroa destructor mites in honey bee colonies. For a accurate judgment of the deaths of bee colonies, in addition to existing requirements bee pathogen control procedures defined by diagnostic Manuals and Code of the OIE, it would be necessary to have the results to prove the qualitative and quantitative presence of the virus in the species Apis mellifera. The introduction and standardization of molecular biological methods in routine diagnosis of viral diseases of bees is justified. It is only a matter of time before the virus infections of bees will be subject to routine diagnostics which controls the health of bee colonies</p>