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Boek Dietary Nitrate and Sports Performance Final Dietary nitrate and sports performance Citation for published version (APA): Jonvik, K. L. (2018). Dietary nitrate and sports performance. Datawyse / Universitaire Pers Maastricht. https://doi.org/10.26481/dis.20180913kj Document status and date: Published: 01/01/2018 DOI: 10.26481/dis.20180913kj Document Version: Publisher's PDF, also known as Version of record 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.umlib.nl/taverne-license 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: 10 Oct. 2021 Dietary nitrate and sports performance The studies presented in this thesis were performed within School of Nutrition and Translational Research in Metabolism. The project was financially supported through a grant from the Dutch Technology Foundation STW (Beet-ing your competitors, project number 12877), and was performed in collaboration between Maastricht University and HAN University of Applied Sciences in a joint consortium with NOC*NSF, Sports Centre Papendal, Kenniscentrum Suiker & Voeding, and NIZO Food Research. Cover design: Hensen Trenning, Karin Lambrechtse and Kristin Lundanes Jonvik Layout: Hensen Trenning, Karin Lambrechtse and Kristin Lundanes Jonvik Printed by: Datawyse, Universitaire Pers Maastricht ISBN 978-94-93019-04-1 © Copyright Kristin Lundanes Jonvik, Maastricht 2018 All rights reserved. No part of this thesis may be reproduced, distributed, or transmitted in any form or by any means, including photocopying, recording, or other electronic or mechanical methods, without the prior written permission of the author or publisher, except in the case of brief quotations embodied in critical reviews and certain other noncommercial uses permitted by copyright law. Dietary nitrate and sports performance DISSERTATION To obtain the degree of Doctor at Maastricht University, on the authority of the Rector Magnificus, Prof. Dr. Rianne M. Letschert in accordance with the decision of the Board of Deans, to be defended in public on Thursday 13 September 2018, at 16:00 hours by Kristin Lundanes Jonvik Born on 24-09-1987 PROMOTOR Prof. dr. L.J.C. van Loon CO-PROMOTOR Dr. L.B. Verdijk ASSESSMENT COMMITTEE Prof. dr. A.M.W.J. Schols (chair) Prof. dr. F. Brouns Prof. dr. A. Jones (University of Exeter, United Kingdom) Prof. dr. R. Meeusen (Vrije Universiteit Brussel, Belgium) Prof. dr. M. Poeze Niet de hoeveelheid overwinningen, maar het aantal foute passes, gemiste kansen en bloedneuzen die daaraan vooraf gingen bepalen hoe succesvol je bent De ‘Wall of failure’ in… Leentje Adres: ‘Meesterlijk Begeleiden’ Postcode: L.E.F. Lerend Experimenterend Filosoferend Heleen van der Wilt Table of contents CHAPTER 1 General introduction 9 CHAPTER 2 Can elite athletes benefit from dietary nitrate 27 supplementation? CHAPTER 3 No effect of acute and 6-day nitrate supplementation 33 on VO2 and time-trial performance in highly-trained cyclists CHAPTER 4 Nitrate-rich vegetables increase plasma nitrate and 47 nitrite concentrations and lower blood pressure in healthy adults CHAPTER 5 Habitual dietary nitrate intake in highly trained athletes 67 CHAPTER 6 Repeated-sprint performance and plasma responses 87 following beetroot juice supplementation do not differ between recreational, competitive and elite sprint athletes CHAPTER 7 The effect of beetroot juice supplementation on 107 dynamic apnea and intermittent sprint performance in elite female water polo players CHAPTER 8 Sucrose but not nitrate ingestion reduces high-intensity 119 exercise-induced gut injury CHAPTER 9 General discussion 137 Summary 157 Valorization 161 Dankwoord 167 Curriculum Vitae List of Publications Chapter General introduction 1 Chapter 1 “Sports nutrition involves the application of eating strategies to promote good health and adaptation to the training program, to recover quickly after each exercise session and to perform optimally during competition” (1). The science of nutrition in relation to sports performance has progressed from empirical studies in the 1960s, on the effects of dietary manipulations, to the direct investigation of the physiological mechanisms of the specific nutritional demands of exercise (2). Within the last decade, there has been an explosion of publications and consensus statements related to sports nutrition. Especially the use of sports supplements to enhance exercise performance has received more attention. Sports supplements represent an ever-growing industry, and the prevalence of supplement use by athletes internationally has been estimated to range between 40-90%, with greater prevalence reported among elite athletes (3). One of the supplements becoming progressively popular amongst athletes is dietary nitrate, due to its proposed beneficial effects on health and performance. Since Larsen et al. in 2007 (4) reported reduced oxygen demands during exercise and thereby more efficient energy use following nitrate supplementation, research on nitrate in athletes has expanded. Even though there is great diversity in the results of studies on nitrate and performance, nitrate-rich beetroot juice is now considered one of the top-5 ergogenic aids (5). As a result, many athletes are currently using beetroot juice as a performance-enhancing supplement. Next to being a popular supplement, nitrate is a component of our habitual diet, and is found in considerable amounts in beetroot and green leafy vegetables, such as rocket salad, spinach and lettuce (6). The beneficial effects of dietary nitrate are related to its ability to increase the bioavailability of nitric oxide in the human body (7). Dietary nitrate and nitric oxide Nitric oxide (NO) represents a key regulator of several physiological functions (Figure 1.1), including vasodilation, muscle contractility, cell cycle regulation, mitochondrial respiration, glucose and calcium homeostasis (8, 9); all of which represent important factors in determining exercise and performance capacity. NO was originally believed to be generated solely through the oxidation of L-arginine – in a reaction catalyzed by nitric oxide synthase (NOS), with nitrite (NO2 ) and nitrate – (NO3 ) considered to be inert by-products of this process. However, in the 1990s a reverse pathway was shown, whereby nitrate and nitrite can be reduced back to NO (10, 11). There is now a general consensus that dietary nitrate ingestion can strongly increase plasma nitrate and nitrite concentrations. 10 General introduction Cell cycle regulation Mitochondrial respiration Ca2+ SR calcium uptake NO Muscle contractility Vasodilation Glut-4 Glucose uptake Figure 1.1: Key regulating physiological functions of nitric oxide (NO). NO is important for the regulation of vasodilation, muscle contractility, cell cycle regulation, mitochondrial respiration, glucose uptake and sarcoplasmic reticulum (SR) calcium uptake. Once ingested, inorganic nitrate from dietary sources is readily converted to nitrite by bacteria present in the oral cavity (12). Nitrite is then swallowed with saliva whereof some is reduced to NO in the acidic environment of the stomach. However, a substantial amount of nitrite enters the circulation, elevating plasma nitrite concentrations. Plasma nitrite is subsequently reduced to NO through several enzymatic reactions in blood and other tissues (8, 12). The nitrate-nitrite-NO pathway (Figure 1.2) may be important as a means to increase NO production under circumstances when NO synthesis by the NOS enzymes is impaired. While the NOS pathway is highly oxygen dependent, the nitrate-nitrite-NO pathway is stimulated under conditions of low pH and low oxygen availability (13). As such, the nitrate- nitrite-NO pathway is likely to be activated when oxygen is a limiting factor, such as under ‘local hypoxia’ in skeletal muscle tissue during high-intensity intermittent type exercise or under ‘systemic hypoxia’ such as at high altitude or under water. 11 Chapter 1 Once produced, NO has a half-life of ~1 s and is quickly oxidized to e.g. nitrite and nitrate (14). Therefore, NO is not sufficiently stable to be measured
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