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University of Southampton Research Repository University of Southampton Research Repository Copyright © and Moral Rights for this thesis and, where applicable, any accompanying data are retained by the author and/or other copyright owners. A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. This thesis and the accompanying data cannot be reproduced or quoted extensively from without first obtaining permission in writing from the copyright holder/s. The content of the thesis and accompanying research data (where applicable) must not be changed in any way or sold commercially in any format or medium without the formal permission of the copyright holder/s. When referring to this thesis and any accompanying data, full bibliographic details must be given, e.g. Thesis: Author (Year of Submission) "Full thesis title", University of Southampton, name of the University Faculty or School or Department, PhD Thesis, pagination. Data: Author (Year) Title. URI [dataset] UNIVERSITY OF SOUTHAMPTON FACULTY OF NATURAL AND ENVIRONMENTAL SCIENCES School of Biological Sciences The Effects of Extremely Low Frequency Electromagnetic Fields on Insects by Sebastian James Shepherd Thesis for the degree of Doctor of Philosophy January, 2018 UNIVERSITY OF SOUTHAMPTON ABSTRACT FACULTY OF NATURAL AND ENVIRONMENTAL SCIENCES School of Biological Sciences Thesis for the degree of Doctor of Philosophy THE EFFECTS OF EXTREMELY LOW FREQUENCY ELECTROMAGNETIC FIELDS ON INSECTS Sebastian James Shepherd Flying insect species are currently in decline, including many species that provide important pollination ecosystem services. Combined exposure to various environmental stressors are associated with insect declines, including land-use change, pesticide use and climate change, but the potential biological and environmental effects of extremely low frequency electromagnetic fields (ELF EMFs) are poorly understood. ELF EMFs are pervasive in the environment, and anthropogenic ELF EMF pollution has increased greatly in recent years. Despite this, little has been done to consider the potential environmental impacts of ELF EMFs. Given that there is evidence that ELF EMFs can have biological effects, it is important to explore these biological stimuli and their potential to affect insects in the environment. Here the biological effects of ELF EMFs on important insect species were investigated, with two-fold aims of both increasing understanding of the biological effects of ELF EMFs, and determining whether field-realisitc levels of ELF EMFs have the potential to cause envrionemental stress to insects. ELF EMF impacts were investigated with the desert locust, as an economically important agricultural pest and a species that provides utility in understanding insect neurophysiology, and the honey bee, as a globally important pollinator and a well known study species for insect cognitive behaviour. Short-term exposure to high levels of ELF EMFs was found to affect neurophysiology, and reduce locomotory function in locusts, as well as increase stress protein levels in bees and locusts, and affect honey bee cognitive behaviour. Acute exposure to ELF EMFs at levels that can be encountered regularly in the environment around man-man sources for ELF EMFs reduced honey bee performance in olfactory learning assays, affected flight behaviour, and affected feeding and flight performance in a semi-field scenario. Further to this, some of these impacts of ELF EMFs on cognitive behaviour and flight were reduced when ELF EMFs were applied in combination with other well-known environmental stressors, neonicotinoid insecticides. These findings give a more detailed indication of some of the physiological effects that may underpin changes in insect locomotory behaviour that occur after short-term exposure. This is the first time that powerline simulating ELF EMFs have been directly measured and applied to insects in the context of considering the ecological effects (and thus using field- realistic exposure levels) of the ELF EMFs, rather than just the biological effects. This is the first indication that short-term and acute ELF EMF exposure can affect insect cognitive behaviour, and these effects have been shown to occur at levels which can be encountered in the field by a globally important pollinator species, the honey bee. This is also the first indication that acute field-realistic ELF EMF exposure can affect insect locomotory behaviour in the environement. This research describes new effects of ELF EMFs on insect biology and establishes that ELF EMFs have a potential to affect insect ecology, such that future ELF EMF understanding must be focused in further exploring mechanisms by with ELF EMFs cause biological effects, as well as the larger scale ecological risk assessment of ELF EMF impacts from powerlines. Table of Contents Table of Contents Table of Contents .................................................................................................................. i Table of Tables .................................................................................................................... ix Table of Figures ................................................................................................................... xi Academic Thesis: Declaration of Authorship ................................................................. xxi Acknowledgements ......................................................................................................... xxiii Definitions and Abbreviations ...................................................................................... xxvii Chapter 1 General Introduction..................................................................................... 1 1.1 Insect ecology, declines, and global impacts ........................................................... 1 1.1.1 Relationship between insects and humans ...................................................... 1 1.1.2 Importance of insects as pests ......................................................................... 1 1.1.3 Importance of insects as pollinators ................................................................ 2 1.1.4 Anthropogenic change ..................................................................................... 3 1.1.5 Electromagnetic fields and anthropogenic change .......................................... 4 1.2 Electromagnetism and its properties ........................................................................ 6 1.2.1 Physical fields .................................................................................................. 6 1.2.2 Electric fields ................................................................................................... 6 1.2.3 Magnetic fields ................................................................................................ 7 1.2.4 Electromagnetic fields ..................................................................................... 8 1.2.4.1 Time-varying EMFs and electromagnetic radiation ................................ 8 1.2.4.2 The electromagnetic spectrum ................................................................. 8 1.2.4.3 Sources of electric and magnetic fields ................................................. 10 1.3 Electromagnetic fields ........................................................................................... 12 1.3.1 Mechanisms by which EMFs interact with physical material ....................... 12 1.3.2 Electromagnetic mechanisms for biological effects ...................................... 14 1.3.3 Potential for biological effects ....................................................................... 14 1.3.4 How EMF exposure leads to biological effects ............................................. 15 1.3.4.1 Biological interactions of electric fields ................................................ 16 1.3.4.2 Biological interactions of magnetic fields ............................................. 17 1.4 Extremely low frequency electromagnetic fields .................................................. 22 1.4.1 Study focus and rationale .............................................................................. 22 i Table of Contents 1.4.2 Sources and scale of ELF EMF emission ..................................................... 23 1.4.3 Examples of ELF EMF effects ...................................................................... 25 1.4.3.1 Molecular examples .............................................................................. 25 1.4.3.2 Physiological effects ............................................................................. 27 1.4.3.3 Behavioural effects ............................................................................... 29 1.4.3.4 Ecological effects of ELF EMF and applications ................................. 30 1.4.3.5 Epidemiological studies ........................................................................ 32 1.4.4 ELF EMF safety, standards, and costs .......................................................... 34 1.4.5 Research needs .............................................................................................. 36 1.5 Studying the effects of ELF EMFs on insects ....................................................... 38 1.5.1 Key attributes of insect research ................................................................... 38 1.5.2 Potential for ecological effects of EMFs on insects ...................................... 39 1.5.2.1 Potential exposure ................................................................................
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