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Human thermoregulation : a synergy between physiology and mathematical modelling Citation for published version (APA): Kingma, B. R. M. (2012). Human thermoregulation : a synergy between physiology and mathematical modelling. Universitaire Pers Maastricht. Document status and date: Published: 01/01/2012 Document Version: Publisher’s PDF, also known as Version of Record (includes final page, issue and volume numbers) Please check the document version of this publication: • A submitted manuscript is the version of the article upon submission and before peer-review. There can be important differences between the submitted version and the official published version of record. People interested in the research are advised to contact the author for the final version of the publication, or visit the DOI to the publisher's website. • The final author version and the galley proof are versions of the publication after peer review. • The final published version features the final layout of the paper including the volume, issue and page numbers. Link to publication General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal. If the publication is distributed under the terms of Article 25fa of the Dutch Copyright Act, indicated by the “Taverne” license above, please follow below link for the End User Agreement: www.tue.nl/taverne Take down policy If you believe that this document breaches copyright please contact us at: [email protected] providing details and we will investigate your claim. Download date: 23. Sep. 2021 Human Thermoregulation A synergy between physiology and mathematical modelling The research presented in this thesis was performed within NUTRIM School for Nutrition, Toxicology and Metabolism which participates in the Graduate School VLAG (Food Technology, Agrobiotechnology, Nutrition and Health Sciences), accred- ited by the Royal Netherlands Academy of Arts and Sciences. Furthermore, the re- search was performed in collaboration with Eindhoven University of Technology, department of Mechanical Engineering. Cover design: Eva Tan & Boris Kingma (Cover picture: http://www.dreamstime.com) Layout: Boris Kingma Printed by: Datawyse / Universitaire Pers Maastricht © Boris Kingma, Maastricht 2011 Universitaire Pers Maastricht ISBN: 978 94 6159 106 7 Human Thermoregulation A synergy between physiology and mathematical modelling PROEFSCHRIFT ter verkrijging van de graad van doctor aan de Universiteit Maastricht op gezag van de Rector Magnificus Prof. mr. G.P.M.F. Mols volgens het besluit van het College van Decanen in het openbaar te verdedigen op vrijdag 27 januari 2012 om 14.00 uur door Boris René Motrona Kingma geboren te Wilrijk (B) op 19 februari 1982 UMP UNIVERSITAIRE PERS MAASTRICHT Promotores Prof. Dr. Ir. W. H. M. Saris Prof. Dr. Ir. A. A. van Steenhoven (TU Eindhoven) Co-Promotores Dr. W. D. van Marken Lichtenbelt Dr. Ir. A. J. H. Frijns (TU Eindhoven) Beoordelingscommissie / Assessment Committee Prof. Dr. G. J. van der Vusse (voorzitter) Dr. N. D. Bouvy Prof. Dr. H. A. M. Daanen (VU Amsterdam / TNO Defence, Security and Safety) Dr. D. Fiala (Ergonsim - Comfort Energy Efficiency / Stuttgart University) Prof. Dr. Ir. R. Peeters Prof. Dr. L. P. A. J. Schrauwen The research described in this thesis was supported by a grant of AgentschapNL (EOS-LT 03001). Financial support for printing this thesis by AgentschapNL, Perimed and CorTemp is greatly acknowl- edged. Table of Contents Chapter 1 General Introduction 9 The thermoneutral zone: implications for metabolic Chapter 2 19 studies Chapter 3 Local control of skin blood flow: the effect of age 41 Increased systolic blood pressure after mild cold and Chapter 4 rewarming: effect of age and cold induced thermo- 55 genesis Incorporating neurophysiological concepts in math- Chapter 5 ematical thermoregulation models 71 Mathematical model of thermal sensation based on Chapter 6 89 the neurophysiology of thermal reception Chapter 7 General discussion 107 Thermophysiological Simulation Model Appendix 117 Eindhoven Maastricht Summary 135 Samenvatting 141 Dankwoord 147 List of Publications 153 Curriculum vitae 157 List of symbols Symbol Description Unit A Surface area [m2] a Distribution or weighting coefficient [-][AU] c Heat capacitance [J kg-1 °C-1] Cs Vasoconstriction [-] D Sign operator for direction of response [-] Dl Vasodilation [-] -1 Neuron fire rate [pulse s ] f͚cl Clothing surface area factor [-] G Gain factor for neuron fire rate [-] h Surface heat transfer coefficient [W m-2 °C-1] -1 hx Counter current heat transfer coefficient [W °C ] H Hypothalamic neuron fire rate pulse s-1 icl Moisture permeability coefficient of clothing [-] Icl Heat resistance of clothing [Clo] k Conductivity [W m-1 °C-1] K Gain factor for dynamic part of neuron fire rate [-] L Length [m] Le Lewis constant [°C Pa-1] m Mass [kg] N Neural regulatory signal for skin blood flow [-] p Probability of obtaining a test-statistic at least as extreme as the one [-] observed pair Air pressure [Pa] P Peripheral afferent neural drive [pulse s-1] -2 q Surface heat transfer [W m ] -3 qm Metabolic heat production [W m ] r Radius [m] -2 -1 Re Skin permeability resistance [W m Pa ] S Steady state neuron fire rate pulse s-1 s Coefficient for steady state neuron fire rate [-] Sh Shivering [W] Sw Sweating [g min-1] t Time [s] T Temperature [°C] -2 -1 Ucl Combined radiative and convective heat transfer coefficient (cloth- [W m °C ] ing) -2 -1 Ue Evaporative heat transfer coefficient (clothing) [W m Pa ] -1 v Speed [m s ] V Volume [m3] Respiration rate [m3 s-1] Vʖ Greek symbols Description Unit β Heat equivalent of blood flow [W m-3] γ Model constants for neurophysiological skin blood flow model [s pulse-1] ∆ Difference [-] ε Emissivity [-] κ Model constants for neurophysiological thermal sensation model [s pulse-1] φ Relative humidity [%][-] ϕ Angle [°] λ Heat of vaporization [J kg-1] µ Proportionality constant for change in blood flow due to change in [°C-1] metabolism ψ View factor [-] ρ Density [kg m-3] σ Stefan Boltzmann constant [W m-2 K-4] ω Perfusion rate [l m-3 s-1] Subscripts and superscripts a Arterial air Air acc Accumulation b Blood bas Basal, baseline c Convective cl Clothing cold Cold e Evaporative eff Effective ex Exhaled exc Excitatory frc Forced H2O Water i Index in Inhaled inh Inhibitory j Index mean Mean m Metabolic max Maximum mix Mixed nat Natural op Operative r Radiative res Respiratory sat Saturated sh Shivering sk Skin sp Specific sr Sector surf Surface sw Sweat sweatgland Sweatgland t Time vap Vaporization w Wall warm Warm x Counter current Abbreviations and acronyms ASHRAE American Society of Heating, Refrigerating and Air-Conditioning Engineers AVA Arterio-venous anastamosis AU Arbitrary unit BAT Brown adipose tissue BMI Body mass index BMR Basal metabolic rate BP Blood pressure BPM Beats per minute BR Bretylium tosylate CIT Cold induced thermogenesis CIZ Core inter-threshold zone CVC Cutaneous vascular conductance DIT Diet induced thermogenesis EE Energy expenditure HR Heart rate HVAC Heating Ventilation Air Conditioning LCT Lower critical temperature LDF Laser Doppler flowmetry MAP Mean arterial blood pressure MSR Mean squared residual NA Noradrenalin (Nor-Epinephrine) NST Non-shivering thermogenesis PMV Predicted Mean Vote PU Perfusion unit RMSR (Square) Root of mean squared residual SBF Skin blood flow SBP Systolic blood pressure SE Standard error of the mean SFT Skinfold thickness SMR Standard metabolic rate SP Set point ThermoSEM Thermophysiological Simulation Model Eindhoven Maastricht TNZ Thermoneutral zone UCT Upper critical temperature USA United States of America VC Vasoconstriction CHAPTER 1 GENERAL INTRODUCTION 9 GENERAL INTRODUCTION Human thermoregulation and the environment Human beings are warm-blooded animals, which means that they are able to main- tain a stable temperature for their vital organs despite large fluctuations in envi- ronmental conditions. Humans do this by carefully balancing heat production and heat loss. If more heat is lost than produced, body temperature will drop. Vice ver- sa, if more heat is produced than lost, body temperature will increase (1). Control of this heat balance is referred to as thermoregulation and is effectively executed by modulating behavior (e.g. changing clothes) and physiological mechanisms (e.g. shivering or sweating). Figure 1: Standard metabolic rate vs. mass in reptiles (ectoterms) and mammals (endo- therms). Modified from ref. (2). Warm-blooded animals are also referred to as endotherms (from Greek, endos: inner, thermos: hot). Endothermy creates opportunities that cannot be exploited by ectotherms (Greek, ectos: outer; e.g. reptiles or amphibians) as long as there is enough fuel to keep the fire burning and enough water to cool the machinery down. For instance, ectotherms rely on environmental conditions to heat up before they can perform vital actions such as foraging or reproducing. Endotherms, in turn, are less dependent on environmental conditions and can therefore live in a wider range of climates, thereby reducing competition for resources and increasing chances of survival. Nevertheless, maintaining a stable body temperature comes at a cost, for 10 GENERAL INTRODUCTION the same body mass endotherms maintain higher energy expenditure than ecto- therms (see Figure 1) (2). Also in cold or hot environments, relatively more resources have to be spent to regulate body temperature than in thermoneutral environments. Therefore, it might not be coincidental that the earliest civilizations developed in climatic regions close to thermoneutrality (see Figure 2). Figure 2: The 21°C isotherm (shown as 70° Fahrenheit) and the sites of ancient civilizations.
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