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Adaptive Homeostasis 4 5 Q1 Kelvin J.A ARTICLE IN PRESS Molecular Aspects of Medicine ■■ (2016) ■■–■■ Contents lists available at ScienceDirect Molecular Aspects of Medicine journal homepage: www.elsevier.com/locate/mam 1 Q2 Review 2 3 Adaptive homeostasis 4 5 Q1 Kelvin J.A. Davies a,b,* 6 a Leonard Davis School of Gerontology of the Ethel Percy Andrus Gerontology Center, The University of Southern California, Los Angeles, CA 7 Q5 90089-0191, USA 8 b Division of Molecular and Computational Biology, Department of Biological Sciences, Dornsife College of Letters, Arts, & Sciences, The Uni- 9 versity of Southern California, Los Angeles, CA 90089-0191, USA 10 11 12 ARTICLE INFO ABSTRACT 13 1514 Article history: Homeostasis is a central pillar of modern Physiology. The term homeostasis was invented 1716 Received 15 April 2016 by Walter Bradford Cannon in an attempt to extend and codify the principle of ‘milieu 1918 Accepted 15 April 2016 intérieur,’ or a constant interior bodily environment, that had previously been postulated 20 Available online 21 by Claude Bernard. Clearly, ‘milieu intérieur’ and homeostasis have served us well for over 2322 a century. Nevertheless, research on signal transduction systems that regulate gene ex- 2524 Keywords: pression, or that cause biochemical alterations to existing enzymes, in response to external 26 Homeostasis 27 and internal stimuli, makes it clear that biological systems are continuously making short- 28 Adaptation 3029 Stress term adaptations both to set-points, and to the range of ‘normal’ capacity. These transient 3231 Hormesis adaptations typically occur in response to relatively mild changes in conditions, to pro- 3433 Nrf2 grams of exercise training, or to sub-toxic, non-damaging levels of chemical agents; thus, 3635 Aging the terms hormesis, heterostasis, and allostasis are not accurate descriptors. Therefore, an 37 operational adjustment to our understanding of homeostasis suggests that the modified 38 term, Adaptive Homeostasis, may be useful especially in studies of stress, toxicology, disease, 39 and aging. Adaptive Homeostasis may be defined as follows: ‘The transient expansion or 40 contraction of the homeostatic range in response to exposure to sub-toxic, non-damaging, 41 signaling molecules or events, or the removal or cessation of such molecules or events.’ 42 © 2016 Published by Elsevier Ltd. 43 44 Contents 45 46 1. Introduction to homeostasis ............................................................................................................................................................................................... 1 47 2. The importance of stress ...................................................................................................................................................................................................... 2 48 3. Heterostasis and allostasis ................................................................................................................................................................................................... 3 49 4. Hormesis .................................................................................................................................................................................................................................... 4 50 5. Adaptive homeostasis ............................................................................................................................................................................................................ 4 51 6. Summary and conclusions ................................................................................................................................................................................................... 5 52 Acknowledgements ................................................................................................................................................................................................................ 6 53 Uncited references .................................................................................................................................................................................................................. 6 54 References .................................................................................................................................................................................................................................. 6 55 56 57 58 Q3 Q4 * Leonard Davis School of Gerontology of the Ethel Percy Andrus 1. Introduction to homeostasis Q6 64 59 Gerontology Center and Division of Molecular and Computational Biology, 65 60 Department of Biological Sciences, Dornsife College of Letters, Arts, & The concept of milieu intérieur, or a constant interior 66 61 Sciences, The University of Southern California, Los Angeles, CA 90089-0191, 62 USA. bodily environment, was developed by the celebrated French 67 63 E-mail address: [email protected]. physiologist Claude Bernard in 1865 (Bernard, 1865). The 68 http://dx.doi.org/10.1016/j.mam.2016.04.007 0098-2997/© 2016 Published by Elsevier Ltd. Please cite this article in press as: Kelvin J.A. Davies, Adaptive homeostasis, Molecular Aspects of Medicine (2016), doi: 10.1016/j.mam.2016.04.007 ARTICLE IN PRESS 2 K.J.A. Davies / Molecular Aspects of Medicine ■■ (2016) ■■–■■ 69 word ‘homeostasis’ was coined by the Harvard Physiolo- 70 gist, Walter Bradford Cannon in 1926 to describe and extend 71 Bernard’s milieu intérieur concept (Cannon, 1926), and popu- 72 larized (in 1932) in his highly successful and persuasive book, 73 The Wisdom of the Body (Cannon, 1932). Cannon com- 74 bined two words from Ancient Greek ο μος (hómos, 75 ”similar”) + ιστημι (histe¯mi, ”standing still”)/stasis (from 76 στα´σις) into a Modern Latin form to invent his term 77 homeostasis. 78 Cannon wrote, “The constant conditions which are main- 79 tained in the body might be termed equilibria. That word, 80 however, has come to have fairly exact meaning as applied 81 to relatively simple physico-chemical states, in closed 82 systems, where known forces are balanced. The coordi- 83 nated physiological processes which maintain most of the 84 steady states in the organism are so complex and so pecu- 85 liar to living beings – involving, as they may, the brain and 86 nerves, the heart, lungs, kidneys and spleen, all working co- 87 operatively – that I have suggested a special designation for 88 these states, homeostasis. The word does not imply some- 89 thing set and immobile, a stagnation. It means a condition Fig. 1. Walter Bradford Cannon (1871–1945). The Harvard Physiologist who Q12 130 coined the term Homeostasis in 1926 to describe and extend Bernard’s 131 90 – a condition which may vary, but which is relatively milieu intérieur concept (Cannon, 1926). Homeostasis was subsequently 132 91 constant.” popularized (in 1932) in Cannon’s highly influential book, The Wisdom of 133 92 Claude Bernard (July 12, 1813–February 10, 1878), who the Body (Cannon, 1932). Image attribution: https://upload.wikimedia.org/ 134 93 is considered by many to have been the “father” of modern wikipedia/commons/7/78/Walter_Bradford_Cannon_1934.jp 135 94 experimental physiology, is quoted as having said that, “The 136 95 laboratory is the temple of the science of medicine.”(Schafer, 96 2009). Working at a time when cells were just beginning 3 The regulating system that determines the homeo- 137 97 to be thought of as the basic structural unit of tissue and static state consists of a number of cooperating 138 98 organ anatomy, Bernard was able to add an entirely new level mechanisms acting simultaneously or successively. 139 99 of functional integration. Bernard concluded that, “The con- 4 Homeostasis does not occur by chance, but is the result 140 100 stancy of the internal environment is the condition for free and of organized self-government (Fig. 1). 141 101 independent life: the mechanism that makes it possible is that 142 102 which assured the maintenance, within the internal environ- According to Arthur C. Guyton’s immensely influential 143 103 ment, of all the conditions necessary for the life of the elements.” Textbook of Medical Physiology (Guyton, 1991), “The term ho- 144 104 (Bernard, 1974a, 1974b; Gross, 1998). An important biog- meostasis is used by physiologists to mean, maintenance of 145 105 raphy of Claude Bernard has been written by Charles Gross nearly constant conditions in the internal environment.” 146 106 (Gross, 1998). The basic idea of homeostasis is shown in Fig. 2 below. 147 107 Prior to gaining his medical degree in 1900, Walter The Y axis of Fig. 2 can be any biological/physiological func- 148 108 Cannon (October 19, 1871–October 1, 1945) was a student tion, such as blood pressure, heart rate, core temperature, 149 109 of Physiologist Henry Pickering Bowditch, who became Dean blood glucose, NAD+/NADH and NADP+/NADPH ratios, su- 150 110 of Harvard’s Medical School. Bowditch, in turn, had studied peroxide dismutase and glutathione peroxidase levels and 151 111 in Paris with Claude Bernard in the late 1860s, after gradu- activities, Proteasome and Lon levels and activities, or the 152 112 ating from Harvard College. It is clearly no accident that capacity and effectiveness of DNA repair systems. The X axis 153 113 Cannon, who in 1906 became Higginson Professor and chair is calibrated by time, whose units can be seconds, minutes, 154 114 of the department of physiology at Harvard Medical School, hours, days, weeks, or even years (if one considers aging).
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