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Sleep in the Arabian Oryx, Oryx Leucoryx

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Sleep in the Arabian Oryx, Oryx Leucoryx Sleep in the Arabian oryx, Oryx leucoryx Joshua Gabriel Davimes 319626 Original published work submitted to the Faculty of Health Sciences, University of the Witwatersrand, Johannesburg, in fulfillment of the requirements for the degree of Doctor of Philosophy Johannesburg, 2018 i DECLARATION I, Joshua Gabriel Davimes declare that this thesis is my own, unaided work. It is being submitted for the degree of Doctor of Philosophy in the University of the Witwatersrand, Johannesburg. It has not been submitted before for any degree or examination at this or any other university. Signature: Date: 20 February 2018 ii DEDICATION “For the pursuit of eternal knowledge and ineffable understanding, one must only dream...” Josh Davimes This work is dedicated to the important people in my life. My parents Claudia and Stephen, my brother Dominic, my Daschunds, Luigi and Franchesca and lastly, my most wondrous companion Camilla. iii ABSTRACT The evolution and function of sleep remains an enigma in modern science. Significant variation can be observed within species and across taxa. Little is understood about how sleep exists, or presents itself in species surviving in extreme conditions. For example, harsh desert ecosystems with notably lower survival rates for resident species. It has been hypothesized that specialized sleep-related behaviours impact thermoregulation to promote survival advantage. The threatened Arabian oryx (Oryx leucoryx), a member of the Cetartiodactyl superorder, is a large desert dwelling ungulate species successfully residing in the arid zones of the Middle East. Arabian Oryx exhibit numerous morphophysiological adaptations and capabilities to cope with their harsh environment. The current thesis provides an examination of the neuroanatomy, physiology and behavioural patterns related to sleep in wild, free-roaming Arabian oryx under natural conditions. It was our objective to determine whether the extreme conditions and climate of the Arabian Desert has led to novelties within the organization of the oryx’s sleep related neuronal architecture, seasonal behaviour patterns and sleep physiology. Using immunohistochemistry and stereology our results indicate the sleep related nuclei in the brain of the oryx exhibit a typical mammalian organizational plan with additional order-specific and novel, species-specific features. Actigraphy revealed that oryx exhibit temporal niche switching patterns seasonally presenting with winter diurnal- and summer nocturnal activity and intermittent patterns during the transitional periods of spring and autumn. Polysomnography results indicate novel sleep patterns between seasons and physiologically confirm activity-based temporal niche switching at a seasonal level. Our work covers the first examination of the Arabian oryx brain sleep centers as well as its year-long activity patterns. Importantly, it is the first examination of physiological sleep in wild mammals within a desert environment. Our results suggest that ambient temperature acts a dominant driving force for the adaptive behavioural and physiological features described. The Arabian oryx is a remarkable species, well adapted to such an extreme environment and its highly plastic survival mechanisms appear unique. Considering the continual desertification of our planet and the predicted consequences of climate change, knowledge of such species and their continued conservation is of vital importance. iv PUBLICATIONS AND PRESENTATIONS ARISING FROM THE WORKS IN THIS THESIS Publications Davimes, J., Alagaili, A., Bennet, N., Mohammed, O., Manger, P., Gravett, N., 2015. Nuclear organization and morphology of the sleep related nuclei in the brain of the Arabian oryx, Oryx leucoryx. Journal of Neurochemistry, 134, p149. Conference proceeding: 25th ISN-APSN-ANS Biennial Meeting, Cairns, Australia, August 23-27th, 2015. ISSN Print: 0022-3042. Author contributions: Conceived and designed the experiments: DJ, MP, and GN. Assisted Surgeries: AA, BN, MO, and MP. Performed the experiments: DJ, MP, and GN. Analyzed the data: DJ, MP, and GN. Contributed reagents/materials/analysis tools: AA, BN, MO, MPR, and GN. Wrote Conference Abstract: DJ. Designed Poster: DJ. Presented at Conference: DJ. Davimes, J.G., Alagaili, A.N., Gravett, N., Bertelsen, M.F., Mohammed, O.B., Ismail, K., Bennett, N.C. and Manger, P.R., 2016. Arabian oryx (Oryx leucoryx) respond to increased ambient temperatures with a seasonal shift in the timing of their daily inactivity patterns. Journal of biological rhythms, 31(4), pp.365-374. https://doi.org/10.1177/0748730416645729 Author contributions: Conceived and designed the experiments: DJG, MPR, and GN. Assisted Surgeries: DJG, GN, AAN, BMF, BNC, MOB, IK, and MPR. Performed the experiments: DJG, GN, AAN, BMF, BNC, MOB, IK, and MPR. Analyzed the data: DJG, MPR, and GN. Contributed reagents/materials/analysis tools: AAN, BMF, BNC, MOB, IK, MPR, and GN. Wrote the paper: DJG, MPR, and GN. Provided significant comments and editing on the paper: DJG, MPR and GN. All authors reviewed the final manuscript prior to journal submission. Rebuttal and Revisions of Manuscript: DJG, MPR, and GN. Davimes, J.G., Alagaili, A.N., Gravett, N., Bertelsen, M.F., Mohammed, O.B., Ismail, K., Bennett, N., Manger, P.R., 2016. A possible non-photic primary zeitgeber seasonally entraining dual-phasing in the Arabian oryx. Journal of Sleep and Sleep Disorders Research. Conference v proceeding: Sleep 2016, Denver, Colorado, United States of America, June 2016. ISSN Print: 0161-8105. Author contributions: Conceived and designed the experiments: DJG, MPR, and GN. Assisted Surgeries: DJG, GN, AAN, BMF, BNC, MOB, IK, and MPR. Performed the experiments: DJG, GN, AAN, BMF, BNC, MOB, IK, and MPR. Analyzed the data: DJG, MPR, and GN. Contributed reagents/materials/analysis tools: AAN, BMF, BNC, MOB, IK, MPR, and GN. Wrote Conference Abstract: DJG. Designed Poster: DJG. Presented at Conference: DJG Davimes, J.G., Alagaili, A.N., Bennett, N.C., Mohammed, O.B., Bhagwandin, A., Manger, P.R. and Gravett, N., 2017. Neurochemical organization and morphology of the sleep related nuclei in the brain of the Arabian oryx, Oryx leucoryx. Journal of Chemical Neuroanatomy, 81, pp.53- 70. https://doi.org/10.1016/j.jchemneu.2017.02.002 Author contributions: Conceived and designed the experiments: DJG, MPR, and GN. Assisted Surgeries: AAN, BNC, MOB, and MPR. Performed the experiments: DJG, BA, MPR, and GN. Analyzed the data: DJG, BA, MPR, and GN. Contributed reagents/materials/analysis tools: AAN, BNC, MOB, BA, MPR, and NG. Wrote the paper: DJG, MPR, and GN. Provided significant comments and editing on the paper: DJG, MPR and GN. All authors reviewed the final manuscript prior to journal submission. Rebuttal and Revisions of Manuscript: DJG, MPR, and GN. Davimes, J.G., Alagaili, A.N., Bertelsen, M.F., Mohammed, O.B., Hemingway, J., Bennett, N.C., Manger, P.R. and Gravett, N., 2017. Temporal niche switching in Arabian oryx (Oryx leucoryx): Seasonal plasticity of 24h activity patterns in a large desert mammal. Physiology and Behavior, 177, pp.148-154. https://doi.org/10.1016/j.physbeh.2017.04.014 Author contributions: Conceived and designed the experiments: DJG, MPR, and GN. Assisted Surgeries: AAN, BNC, MOB, and MPR. Performed the experiments: AAN, BNC, MOB, MPR, and GN. Analyzed the data: DJG, HJ, MPR, and GN. Contributed reagents/materials/analysis tools: AAN, BNC, HJ, MOB, MPR, and GN. Wrote the paper: DJG, MPR, and GN. Provided significant comments and editing on the paper: DJG, HJ, MPR, and GN. All authors reviewed the final manuscript prior to journal submission. Rebuttal and Revisions of Manuscript: DJG, MPR, and GN. vi Davimes, J.G., Alagaili, A.N., Bhagwandin, A., Bertelsen, M.F., Mohammed, O.B., Bennett, N.C., Manger, P.R. and Gravett, N., 2018. Seasonal variations in sleep of free-ranging Arabian oryx (Oryx leucoryx) under natural hyper-arid conditions. Sleep. In Press. Author contributions: Conceived and designed the experiments: DJG, MPR, and GN. Assisted Surgeries: AAN, BNC, MOB, BA, and MPR. Performed the experiments: AAN, BNC, MOB, MPR, BA, and GN. Analyzed the data: DJG, MPR, AB, and GN. Contributed reagents/materials/analysis tools: AAN, BNC, MOB, MPR, AB, and GN. Wrote the paper: DJG, MPR, and GN. Provided significant comments and editing on the paper: DJG, MPR, and GN. All authors reviewed the final manuscript prior to journal submission. Rebuttal and Revisions of Manuscript: DJG, MPR, and GN. Conference Presentations Joshua G. Davimes, Abdulaziz N. Alagaili, Mads F Bertelsen, Nigel C. Bennett, Osama B. Mohammed, Paul R. Manger, Nadine Gravett. Title: Seasonal effects on sleep architecture by means of actigraphy in free-roaming Arabian Oryx, Oryx leucoryx. 12th International Society of Neuroscientists of Africa (SONA) Conference- Durban, South Africa. March 26th - 30th 2015. Joshua G. Davimes, Abdulaziz N. Alagaili, Nigel C. Bennett, Osama B. Mohammed, Paul R. Manger, Nadine Gravett. Title: Nuclear organization and morphology of the sleep related nuclei in the brain of the Arabian Oryx, Oryx leucoryx. 43rd Annual Conference of the Anatomical Society of Southern Africa- Parys, South Africa. May 9th - 13th 2015. Joshua G. Davimes, Aziz N. Alagaili, Nigel C. Bennett, Osama B. Mohammed, Paul R. Manger, Nadine Gravett. Title: Nuclear organization and morphology of the sleep related nuclei in the brain of the Arabian Oryx, Oryx leucoryx. 9th IBRO world congress- Rio de Janeiro, Brazil. July 7th - 11th 2015. Joshua G. Davimes, Abdulaziz N. Alagaili, Nigel C.
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