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THE EFFECTS of NATURAL and ANTHROPOGENIC FACTORS on the PHYSIOLOGICAL STATE of INSECTS IRJA KIVIMÄGI a Thesis for Applying

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THE EFFECTS of NATURAL and ANTHROPOGENIC FACTORS on the PHYSIOLOGICAL STATE of INSECTS IRJA KIVIMÄGI a Thesis for Applying THE EFFECTS OF NATURAL AND ANTHROPOGENIC FACTORS ON THE PHYSIOLOGICAL STATE OF INSECTS LOODUSLIKE JA ANTROPOGEENSETE FAKTORITE MÕJU PUTUKATE FÜSIOLOOGILISELE SEISUNDILE IRJA KIVIMÄGI A thesis for applying for degree of Doctor of Philosophy in Entomology Väitekiri filosoofiadoktori kraadi taotlemiseks entomoloogia erialal Tartu 2012 EESTI MAAÜLIKOOL ESTONIAN UNIVERSITY OF LIFE SCIENCES THE EFFECTS OF NATURAL AND ANTHROPOGENIC FACTORS ON THE PHYSIOLOGICAL STATE OF INSECTS LOODUSLIKE JA ANTROPOGEENSETE FAKTORITE MÕJU PUTUKATE FÜSIOLOOGILISELE SEISUNDILE IRJA KIVIMÄGI A thesis for applying for degree of Doctor of Philosophy in Entomology Väitekiri filosoofiadoktori kraadi taotlemiseks entomoloogia erialal Tartu 2012 Institute of Agricultural and Environmental Sciences Estonian University of Life Sciences According to verdict No 119 of October 9, 2012, the Doctoral Committee of the Agricultural and Natural Sciences of the Estonian University of Life Sciences has accepted this for thesis for the defence of degree of Doctor of Philosophy in Entomology. Opponent: Prof. Habil. Vincais Būda Institute of Ecology Vilnius University Lithuania Supervisors: Dr. Luule Metspalu Institute of Agricultural and Environmental Sciences Estonian University of Life Sciences Dr. Angela Ploomi Institute of Agricultural and Environmental Sciences Estonian University of Life Sciences Defence of the thesis: Estonian University of Life Sciences, Karl Ernst von Baer House, Veski 4, Tartu, on November 15, 2012 at 10.00. The English language was edited by Dr. Ingrid H. Williams and Dr. Eha Kruus, and the Estonian by Dr. Luule Metspalu. Publication of this thesis is granted by the Estonian University of Life Sciences and by the Doctoral School of Earth Sciences and Ecology created under the auspices of European Social Fund. © Irja Kivimägi, 2012 ISBN 978-9949-484-48-5 (print) ISBN 978-9949-484-49-2 (pdf) CONTENTS LIST OF ORIGINAL PUBLICATIONS ......................................... 7 ABBREVIATIONS ........................................................................... 9 1. INTRODUCTION ................................................................. 10 2. REVIEW OF THE LITERATURE .......................................... 13 2.1. Respiration measured by CO2 release ............................... 13 2.2. The pattern of O2 uptake ................................................. 14 2.3. Muscular ventilation (pumping) ....................................... 16 2.4. The origin and functions of discontinuous gas exchange .. 17 2.5. Respiratory failures as indicators of chemical stress ........... 20 2.6. Effects of food plants on the development and diapause in insects .......................................................................... 21 3. AIMS OF THE STUDY .......................................................... 23 4. MATERIALS AND METHODS ............................................. 24 4.1. Insects .............................................................................. 24 4.2. Treatments ....................................................................... 24 4.2.1. Pieris brassicae (I) .................................................... 24 4.2.2. Pterostichus niger (II) ............................................... 24 4.2.3. Platynus assimilis (III-IV) ........................................ 25 4.2.4. Mamestra brassicae (V) ............................................ 26 4.3. Measuring methods .......................................................... 27 4.3.1. Coulometric respirometry (I, III, V) ....................... 27 4.3.2. Flow-through CO2 respirometry (I-V) ................... 29 4.3.3. Infrared probe actography (I-IV) ............................ 30 4.3.4. Calorimetry (V) ..................................................... 31 4.4. Data acquisition and statistical analysis ............................ 32 5. RESULTS ................................................................................. 33 5.1. Oxygen convective uptakes in gas exchange cycles in early diapause pupae of Pieris brassicae (I) ................................. 33 5.2. Effects of humidity conditions on the gas exchange patterns on Pterosticus niger (II) ..................................................... 35 5.2.1. Discontinuous gas exchange ................................... 35 5.2.2. Cyclic gas exchange ................................................ 35 5.3. The gas exchange patterns and the effects of sublethal doses of pyrethroid on gas exchange in Playunus assimilis (III, IV) ... 37 5.4. Effects of larval food plants on diapause of the cabbage moth, Mamestra brassicae (III) .................................................... 39 6. DISCUSSION ......................................................................... 42 6.1. Cyclic gas exchange with Passive Suction Inspiration (PSI) in diapausing pupae of Pieris brassicae (I) .............................. 42 6.2. Gas exchange in the carabid beetle Pterostichus niger in low and high humidity conditions (II) .................................... 44 6.3. Interaction between circulation and gas exchange ............. 46 6.4. Gas exchange in the carabid beetle Platynus assimilis before and after treatment with sublethal doses of pyrethroid ..... 47 6.5. The influence of food plants on development and on dor- mancy of Mamestra brassicae (V) ...................................... 49 7. CONCLUSIONS .................................................................... 52 REFERENCES ............................................................................... 54 SUMMARY IN ESTONIAN .......................................................... 67 ACKNOWLEDGEMENTS ........................................................... 71 PUBLICATIONS ........................................................................... 73 CURRICULUM VITAE .............................................................. 124 ELULOOKIRJELDUS ................................................................. 128 LIST OF PUBLICATIONS .......................................................... 131 6 LIST OF ORIGINAL PUBLICATIONS The present thesis is a summary of the following research papers, which are referred to by their Roman numerals in the text. I Jõgar K., Kuusik A., Ploomi A., Metspalu L., Williams I.H., Hiiesaar K., Kivimägi I., Mänd M., Tasa T., Luik A. 2011. Oxygen convec- tive uptakes in gas exchange cycles in early diapause pupae of Pieris brassicae. The Journal of Experimental Biology, 214, 2816 – 2822. II Kivimägi I., Kuusik A., Jõgar K., Ploomi A., Williams I.H., Met- spalu L., Hiiesaar K., Sibul I., Mänd M., Luik A. 2011. Gas exchange patterns of Pterostichus niger (Carabidae) in dry and moist air. Physi- ological Entomology, 36, 62 – 67. III Kivimägi I., Ploomi A., Metspalu L., Švilponis E., Jõgar K., Hiie- saar K., Luik A., Sibul I., Kuusik A. 2009. Physiology of a carabid beetle Platynus assimilis. Agronomy Research, 7 (special issue I), 328 – 334. IV Kivimägi I., Kuusik A., Ploomi A., Metspalu L. Jõgar K., Williams I.H., Mänd M., Sibul I., Hiiesaar K., Luik A. Gas exchange pat- terns in Platynus assimilis (Coleoptera, Carabidae): respiratory failure induced by a pyrethroid. European Journal of Entomology. (Accepted August 17th 2012). V Metspalu L., Kruus E., Jõgar K., Kuusik A., Williams I.H., Veromann E., Luik A., Ploomi A., Hiiesaar K., Kivimägi I., Mänd M. Larval food plants can regulate the cabbage moth, Mamestra brassicae L. population. Bulletin of Insectology. (Submitted). The papers are reproduced by kind permission of the corresponding jour- nal or publisher. 7 Authors’ contribution to each article Idea and Analysis Manuscript Paper Data collection study design of data preparation I KJ, IK, AP, LM, IK, KJ, AK, TT, IK, KJ KJ, IK, AP, IHW, AK, IHW, AK, KH, MM, AP, LM KH, LM, TT, AL AL, TT II IK, KJ, AP, KH, AK, IK, AK, AP, LM, IK, KJ IK, AK, KJ, AP, LM, LM, MM, AL KJ IHW, IS, KH, AL III IK, KJ, AP, KH, AK, IK, KJ, AK, AP, IK IK, AP, LM, EŠ, KJ, LM, AL AL, KH, LM IS, KH, AK AL IV IK, LM, AP, AK, KJ, IK, AP, KJ, AK, IK IK, AK, AP, LM, KH, AL, MM LM IHW, KH, KJ, IS, AL V LM, KJ, IK, EŠ, AP, LM, KJ, IK, AP, LM, IK LM, EŠ, KJ, IK, EV, AL, KH, MM, AK KH AP, IHW, KH, MM, AK, AL AK – Aare Kuusik, AL – Anne Luik, AP - Angela Ploomi, EŠ – Eha (Švilponis) Kruus, EV – Eve Veromann, IK – Irja Kivimägi, IS – Ivar Sibul, IHW – Ingrid H. Williams, KH – Külli Hiiesaar, KJ – Katrin Jõgar, LM – Luule Metspalu, MM – Marika Mänd, TT – Tea Tasa 8 ABBREVIATIONS a.i. active ingredient CGE cyclic gas exchange CO2 carbon dioxide C-phase closed phase DGE discontinuous gas exchange F-phase flutter phase H2O water IR infrared IRGA infrared gas analyser L litre O2 oxygen O-phase open phase PSI passive suction inspiration PSV passive suction ventilation RH relative humidity RMR resting metabolic rate SMR standard metabolic rate VCO2 rate of CO2 emission VO2 rate of O2 emission WLR water loss rate µV m W rate of heat flow 9 1. INTRODUCTION The physiological state of the insect is influenced throughout by environ- mental factors, such as temperature and humidity, by anthropogenic toxi- cants such as pesticides and by insect food plants. Insects respond to these factors with adequate changes in physiological state. To describe changes in insect physiological state it is necessary to understand responses to any influencing factors in the insect. Changes of respiratory physiology show the level of activity and vitality of the insect. The main differ- ence between inactive and active insects reflects metabolic levels and gas exchange patterns.
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