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In the Beat of a Heart: Life, Energy, and the Unity of Nature INDEX A Adenosine triphosphate, 32, 118, 129, 140 Adrenaline, 34 Aitken, A. C., 250 Alaska, 9–10 Alcohol, 62 Allen, Drew, 228, 245 Allen, Joel, 41–42 Allometry definition, 58 ecological studies, 96 mathematics, 59, 82 patterns in body structure and function, 59, 78–79 purpose, 58, 76 See also Power laws Alveolus, 30 Anatomy environment and, 41–44 evolutionary theory, 11–12 Anderson, Bill, 29 Archaea, 246 Aristotle, 1, 6, 14, 32 Autotrophs, 174–175 B Bacteria, 246 Banavar, Jayanth, 114 Banks, Joseph, 206 Bates, Henry, 220 Bell, Graham, 197 Benedict, Francis, 62–64 Bergmann, Carl, 41, 43, 44 Bertalanffy, Ludwig von, 123–125, 243 Bignoniaciae, 145–146 Biodiversity body size and, 93–94 climate and, 206–207, 211–212, 213–216, 225, 227–229 determinants of, 171–174, 185–186, 229 distribution by altitude, 213, 218 ecosystem surveys, 145–153 energy supply and, 214–215 environmental change and, 189–190 food chain distribution, 174–177 fractal model, 181–182, 187–189 generalizability of rules regarding, 235–240 global distribution patterns, 217–218 historical factors, 219–224, 233

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In the Beat of a Heart: Life, Energy, and the Unity of Nature home-range size and, 225–227 Humboldt’s research, 212–213 limits of current knowledge, 245–248 metabolism and, 202–203, 227–229 mid-domain effect, 217–219 neutral ecology model, 196–197, 198–199, 219, 237 niche model, 177–179, 181, 186–187, 202 power laws, 187–188 predator effects, 190 probability modeling, 193–194 resource competition and sharing models, 177, 178, 179, 180–186 size of ecological zone and, 224–227 size of living things and, 233 unifying theory, 202–203 universal biodiversity number, 195–196 Biological scaling, 3, 239–240 biomechanical correlations, 80 four-dimensional conceptualization, 85, 113–114 fractal geometry and, 90, 127 network modeling, 102, 108, 110–111 patterns, 78–79 in single-cell organisms, 116–119 Biology, science of consistency of rules in, 43–44 generalization in, 232–233, 235–240 government support, 87–88 historical development, 5–6, 11–12 information in, 244–245 limits of current knowledge, 245–248 mathematics and, 2, 18–19, 23–25, 88, 242–243 pattern recognition in, 235–236 physics and, 18, 97–98, 231–232, 234–235, 237 Thompson’s mathematical studies, 12–23 vitalist philosophy, 2, 15 Biomechanics, 80, 81–82 Blood supply, 90–92, 103, 114–115 animal size and, 107–108 fluid dynamics, 108–110 fractal geometry, 106–107, 108, 115 Blum, Jacob, 85 Body size allometric measurement, 58–61 biodiversity and, 93–94, 233 bone structure and, 39–40 cell biology and, 72, 73 drug dosing considerations, 69 ecological research, 94–95 ecological resource competition and, 181–182 energy metabolism and, 38–40, 41, 44–50, 57, 73–78, 134 environment and, 41–44 evolutionary patterns, 42 in food chains, 176 forest growth patterns, 153–154, 167–168, 169 harvesting and, 125–126 home-range size and, 160–161 individual development and, 123, 125–126, 128–129 life span and, 88–89, 142–143 limits of tree growth, 163–164 niche model of biodiversity and, 181, 182 organ size and, 73 population density and, 154–162 prospects for cross-species model of metabolism, 83–85 reproductive biology and, 131 resource distribution network design and, 107–108 risk of extinction and, 95–96 significance of, in biological research, 3–4, 39 Boecklen, William, 184 Bolivar, Simon, 209 Boltzmann, Ludwig, 3 Boltzmann factor, 119, 120 Bone structure biomechanics, 81–82 body size and, 39–40

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In the Beat of a Heart: Life, Energy, and the Unity of Nature Bonpland, Aimé, 208, 212 Botany, 93 Boyle, Brad, 148, 149, 151 Bradfield, Hannah Stillman, 49–50 Brain body weight and, 73 metabolism, 31 Brody, Samuel, 49–50, 55, 61 Brown, Jim, 93–94, 96, 98–99, 101–103, 107–108, 111–112, 115–116, 119, 132, 155–156, 186, 227, 236, 239, 244 C Calder, William, 94, 161 Calorie consumption body mass and, 57 evolution of scientific study, 35 measurement, 36 rate, 33 Calorimetry, 35–36 Cancer, 128 Capillaries, 31, 90, 107–108 Carbon, 129, 130 Carbone, Chris, 158, 159, 161, 242 Carlos IV, King, 208–209 Carnivores, 157–159 Cell biology aging effects, 138 animal size and, 72, 73, 74 biological resource networks and, 103 energy metabolism, 34, 71, 72–73, 104, 110, 134 free-radical damage, 137–138 genome size and, 244 growth patterns, 127–128 human metabolism, 30–32 membrane structure, 115 single-cell organisms, 116–119 surface area, 40 See also Mitochondria Chaos theory, 91–92 Chase, Jonathan, 202 Chemistry, 1–2, 15 of individual development, 129 metabolic, 119–120 mitochondrial, 140 Climate change, 42, 164–165, 238 Cold-blooded animals. See Ectotherms Colwell, Robert, 216–218 Complexity theory, 91–92, 243 Conservation efforts, 94–96, 159–160, 238 Costarelli, Vasiliki, 29, 32, 35 Crick, Francis, 25, 234 D da Vinci, Leonardo, 109 Damuth, John, 114, 157 Darwin, Charles, 6, 161, 211, 222 Dayan, Tamar, 43, 44 Development, individual animal size and, 128–129 biological diversity in, 121–122 chemical aspects, 129 determinants of, 129–130 forest growth patterns, 153–154 growth rate, 122–128 limits of tree growth, 163–164 population growth patterns and, 154–162, 167–168, 169 resource allocation in, 168–169 Diamond, Jared, 182–183, 185 Diet and nutrition evolution of scientific study, 35–37 food chains, 174–177 food webs, 200–202 growth and, 129 home-range size and, 161 life span and, 139 metabolism and, 34, 78–79 population growth patterns, 158–162 2,4-Dinitrophenol, 140 Dosimetry, 61–62, 68–70 Drury, Maureen, 10 DuBois, Delafield, 48–49

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In the Beat of a Heart: Life, Energy, and the Unity of Nature DuBois, Eugene, 47, 48–49, 50, 54, 61, 62 Dürer, Albrecht, 16 Dyson, Freeman, 235 E Ecological studies ecosystem metabolism, 164–166 energy concepts in, 93, 96, 97 food web interactions, 200–202 forest growth patterns, 153–157 macroecological approach, 98–102 metabolic ecology, 202–203, 244–245 modeling ecosystem change, 199–201 physics concepts in, 96–100 prospects for unifying theory, 244–245 random change in environments, 190–197 scientific method, 183–185, 197–199 significance of animal size, 93–96 similarities among forests, 162–163, 166–168, 170, 205–206 species migration, 191, 192, 194 See also Biodiversity Ecstasy, 140 Ectotherms, 76–77 Edinburgh Academy, 4, 5 Elastic similarity, 81–82 Elements of Physical Biology, 97 Elephants, 67–70, 160, 251 Emerson, Brent, 216 Endotherms, 41 Energy as basis for unifying theory in biology, 243–245 biodiversity and, 214–215 biological, 3–4, 32–33 distribution in ecosystem development, 155 distribution in food chains, 175–176 ecological studies, 93, 96, 97 extra-metabolic consumption patterns, fertility and, 132–133 food, 36 human metabolism, 32–35 rate of evolution and, 228 social distribution model, 133 See also Metabolic rate Energy equivalence rule, 157–158 Enquist, Brian, 92–93, 96, 101–103, 107–108, 111–112, 115–116, 126, 127, 146–147, 150, 151, 155–156, 162, 166, 236, 240, 241–242, 245 Environment anatomy and, 41–44 climate change, 164–165 individual and population growth patterns, 158–162 metabolism and, 76 See also Ecological studies Eukaryotes, 117 Evolutionary theory animal size, 42 biological distribution networks, 118 climate and, 227–228, 238 energy model, 98 environmental diversity and rate of evolution, 216 life span and, 141–143 neutral model, 196 optimality modeling and, 105–106 resource competition in, 178, 180–181 Thompson’s research and beliefs, 11–12, 19, 20–21 Extinction body size and risk of, 95–96 home-range size and risk of, 225–226 patterns, 199–200 probability model of biodiversity change, 193–194 F Fat, dietary, 34 Feldman, Henry, 83

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In the Beat of a Heart: Life, Energy, and the Unity of Nature Ferguson, Steven, 42 Fisheries research, 9–10 Fluid dynamics, 108–110, 168–169 FLUXNET, 165 Food webs, 174–177, 200–201 Foraminifera, 12–13, 212, 228, 245 Forest Dynamics Plot, 193 Forster, Georg, 208 Foster, Michael, 13–14 Four dimensional modeling, 85, 113–114 Fractal geometry biological resource networks, 106–107, 108, 111–112, 116, 236, 242 blood supply modeling, 90–92, 115 conceptual basis, 89–90 implications for biological research, 90 scaling of metabolic rate, 127 species diversity and, 181–182, 187–189 Free radicals, 137–138, 140, 141, 142 Fuhrman, Frederick, 71–72 G Galileo, 39–40, 91 Gamelan, 143–144 Gamgee, Arthur, 7–12 Gamgee, Fanny, 5 Gamgee, Joseph, 5 General Systems Theory, 124–125 Genetic science chemistry, 129 information theory, 244 limits of current knowledge, 245–246 temperature effects on mutation, 227–228 Thompson and, 19–20 Gentry, Alwyn, 145–146 Gillooly, Jamie, 119 Ginzburg, Lev, 236 Gittleman, John, 158, 242 Glossary of Greek Birds, The, 14 Glossary of Greek Fishes, A, 22 Gould, Stephen Jay, 24, 105 Gravity, 74 Growth individual, 122–123, 125–127 population, 161–162 Gulliver’s Travels, 57–58 H Haber, Fritz, 130 Haldane, J. B. S., 40, 113 Harman, Denham, 137–138 Harris, Arthur, 63 Hartog, Marcus, 250 Heart rate, 134 Hemoglobin, 30–31 Heterotrophs, 174–175 Heusner, Alfred, 82–83 Hindu religion, 32 Historium Animalium, 6, 14 Homeotherms, 41 Home-range size and patterns, 160–161, 217, 225–227 Horn, Henry, 182 Hubbell, Steve, 167, 192, 193, 194–196, 245 Humboldt, Friedrich Heinrich Alexander von, 207–213, 231, 240, 250 Humboldt current, 206 Hutchinson, G. Evelyn, 171, 172, 173–174, 175, 176, 182, 214, 219, 250 Huxley, Aldous, 60 Huxley, Andrew, 60 Huxley, Julian, 60–61 Huxley, Thomas Henry, 60, 64 I Ideas for a Physiognomy of Plants, 211 Information, role in biology, 244–245 Invasive species, 191

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In the Beat of a Heart: Life, Energy, and the Unity of Nature J Jablonski, David, 228 Joule, 33 K Kant, Immanuel, 231–232 Keller, Evelyn Fox, 24 Kerkhoff, Drew, 150, 151, 242 Keystone species, 200–201 Kleiber, Max, 51–53, 55–58, 61, 64, 71, 73–75, 77, 79, 82 Kolm, Niclas, 216 Krebs, Hans, 71 L Laplace, Pierre Simon, 35 Lariviere, Serge, 42 Lavoisier, Antoine, 33, 35, 241 Lees, David, 218 Leibold, Matthew, 202 Leptin, 34 Lewontin, Richard, 105 Life span biological determinants of, 88–89 body size and, 88–89, 142–143 cell biology and, 140 diet and, 139 evolutionary factors, 141–143 free-radical damage and, 137–138 growth patterns, 122–128 metabolism and, 88, 133–137, 139–140, 142–143 reproductive biology and, 141–142 Lindemann, Ray, 175, 176–177 Lotka, Alfred, 96–97, 124 LSD, 67–69 M MacArthur, Robert, 179–181, 184–185, 193–194, 233, 234 Macroecology, 98–102 Madagascar, 218 Malthus, Thomas, 161 Marine biology chemistry, 130 metabolism of sea creatures, 76 mid-domain effect, 218 Thompson’s studies, 7–10, 12–13 Maritan, Amos, 114 Marsupials, 139 Martin, Arthur, 71–72 Mathematics allometric calculations, 59 biology and, 2, 18–19, 23–25, 88, 242–243 ecological modeling, 197–198 theoretical ecology, 98 Thompson’s biological studies, 12–23 Maurer, Brian, 100 Maxwell, James Clerk, 2 May, Robert, 182, 234–235, 237–238, 243 Mayr, Ernst, 44, 234 McMahon, Thomas, 80–82, 83 McNab, Brian, 142–143 Medawar, Peter, 11, 23, 26 Medical science dosimetry, 61, 68–70 use of body surface area measurements, 61–63 Meditation, 34–35 Meeh, Karl, 45, 47 Meiri, Shai, 43, 44 Mencken, H. L., 135 Metabolic rate allometric calculations, 59 animal size and, 38–40, 41, 44–50, 57, 73–78, 83–85 basal rate, 47–48 as basis for unifying theory in biology, 243–245 biodiversity and, 202–203, 227–229, 244–245 biological networks and, 102–114, 115–116

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In the Beat of a Heart: Life, Energy, and the Unity of Nature biomechanics and, 81–82 blood supply modeling and, 91–92 body mass and, 56–58, 63, 64–65 body temperature regulation and, 70, 119–120 calorie consumption, 33, 36 cell biology, 71, 72–73, 74, 104, 110, 115, 134 change in, 33, 34 comparisons between species, 63, 75–77, 82–83 diurnal variation, 82 drug dosing considerations, 69 earliest research, 35–37, 70 of ecological communities, 175 energy equivalence rule, 157–158 environment and, 76 fractal scaling patterns, 90, 236 Harris–Benedict equations, 62–63 home-range size and, 160–161 individual growth patterns and, 124–125 life span and, 88, 133–137, 139–140, 142–143 metabolic ecology, 202–203, 244–245 plant biology, 100–102 population growth patterns and, 155–159 regulation, 34–35, 70–73 reproductive biology and, 131–133 respiratory process, 30–32 resting rate, 35, 48 of single-cell organisms, 117–119 surface area law, 44–50, 53–55, 61–62, 64, 74–75 tissue and organ studies, 70–72 Microbes, 246–248 Mid-domain effect, 217–219 Mitochondria chemistry, 140 concentration, 72 life span and, 140 structure and function, 31–32, 40, 104, 118 uncoupling, 140 Morphology, 3 Moses, Melanie, 132 Muscle biomechanics, 81–82 Musical instruments, 143–144, 182 Musth, 67–68 N Napoleon, 212 Natural selection, 11, 12, 19, 20, 98, 221–222. See also Evolutionary theory Networks, biological blood system modeling, 103, 106–111 body size and, 107–108, 110–111 cell biology and, 103–104 conceptual basis, 102, 126, 236 current scientific thinking, 115–116 fractal geometry, 106–107 metabolic theory, 155–157, 236, 242 minimal flow models, 114–115 optimality modeling, 104–106 plant biology, 112–114 response to, 111–112 single cell organisms, 116–119 Neurophysiology of metabolism, 34–35 Neutral ecology, 196–197, 198–199, 219, 237 Neutral evolution, 196 Newborns, 72–73, 140 Niches, ecological, 177–179, 181, 186–190, 202 Niklas, Karl, 126–127, 162, 168–169 Nitrogen, 129, 130 ecosystem metabolism, 165–166 O Obesity, 34, 73 Olff, Han, 187, 188–189 On Growth and Form, 3, 15, 16–18, 21–23, 24, 25–26, 106, 112, 123, 231 Optimality modeling, 104–106

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In the Beat of a Heart: Life, Energy, and the Unity of Nature Orozco, Gabriel, 26 Otto, Frei, 26 P Parasites, 157 Pearl, Raymond, 135–137 Pearson, Karl, 135 Penis bones, 42 Petrie, George, 13 Phosphorus, 129, 130 ecosystem metabolism, 165–166 Physics, science of biological resource network modeling, 108–112 biology and, 2, 15–16, 18, 97–98, 231–232, 234–235, 237 biomechanics, 80, 81–82 government support, 87 scaling problems in, 88–89 scientific method, 198, 234–235 Pierce, Chester, 67–68 Pither, Jason, 150–151 Plant biology distribution networks, 112–114 diversity surveys, 145–153 ecosystem metabolism, 164–166 forest growth patterns, 153–157, 169, 192–193 individual growth patterns, 78, 126–127, 169 limits of tree growth, 163–164 metabolism studies, 100–102 resource allocation in individual development, 168–169 similarities among forests, 162–163, 166–168, 170, 205–206 tropical forest ecology, 206–207 Power laws biodiversity and size of ecological zone, 225 definition, 59 fractal geometry and, 89–90 model of biodiversity, 187–188 significance of, 243 size of individuals and population density, 154–162 See also Allometry Principles of Botany and of Vegetable Physiology, The, 206 Private Life of the Gannets, The, 60 Problems of Relative Growth, 60 Prokaryotes, 246 Protein, 129 in diet, 36, 37 Purvis, Andy, 233 Q Quantum physics, 88–89 Quarter-power laws, 78–79 R Rameaux, Jean-Francois, 41 Redfield, Alfred, 130 Relativity, theory of, 88–89 Reproductive biology birth rate, 132 growth and, 130–131 life span and, 141–142 metabolism and, 131–133 of microbes, 247 population patterns, 132, 161–162 species diversity, 121, 122 species size and, 131 Respiratory process efficiency, 104 metabolism, 30–32, 33 in plants, 101, 164–165 in single-cell organisms, 118 Richet, Charles, 46 Rinaldo, Andrea, 114 Ritchie, Mark, 187, 188–189 Rosenzweig, Michael, 225–226 Rubner, Max, 36, 37, 38–39, 45, 46, 51, 54, 64, 70, 134, 241

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In the Beat of a Heart: Life, Energy, and the Unity of Nature S Santa Fe Institute, 91–92 Sarrus, Pierre, 41 Scaling botany studies, 93 physics studies, 88–89 plant metabolism, 100–102 See also Biological scaling Seals, 9–10 Self-similarity, 89–90 Self-thinning, 154–155 Simberloff, Daniel, 183–184, 185 Skillets, 182 Slobodkin, Lawrence, 234, 243 Speakman, John, 139, 140 Sponges, 13 Statistical research, 135–136 Stefansson, Vilhjalmur, 49 Stevenson, Robert Louis, 4 Superconducting Supercollider, 87 Surface area law, 44–50, 53–55, 61–62, 64, 74–75, 85 Surface tension, 15 Svensson, Nate, 148–150, 167–168 T Taoism, 32 Temperance movement, 62 Temperature biodiversity and, 227–229 body size and, 41–44, 45–46, 57 comparisons between species, 75–76 ecosystem metabolism and, 165–166 metabolism and, 38–39, 70, 119–120 population patterns and, 162 rate of evolution and, 227–228 Terborgh, John, 225–226 Theory of Transformations, 19 Thermodynamics, 36, 97 Thomas, Chris, 238 Thomas, Warren, 68 Thompson, D’Arcy Wentworth, 1–3, 40, 60–61, 106, 112, 122, 123, 231, 249–251 achievements and contributions, 23–27 early biological research, 7–12 early life and education, 5–6 On Growth and Form, 3, 16–18, 21–22, 25–26, 106, 123 last years, 22–23 multidisciplinary studies, 12–21 personal qualities, 6–7, 23 translation of Aristotle, 1, 6, 14 Thompson, D’Arcy Wentworth (father), 4–5 Thompson, John Skelton, 4 Tilman, David, 186, 243 Tropical Nature, 221 Tusko, 67–69 Tyrannosaurus rex, 80 U Unifying theory biodiversity dynamics, 202–203 biological energy, 93 in biology, 232–233, 235–240, 243–245 ecological niche sharing, 181 function of, 240–243 General Systems Theory, 124 goals of, 2–3 of metabolism and body size, 83–85 physical approaches to biology, 97–98 prospects for, 251–252 Thompson’s multidisciplinary studies, 12–17, 23 Universal biodiversity number, 195–196 V Vegetarians, 63 Vital Energetics, 63

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In the Beat of a Heart: Life, Energy, and the Unity of Nature Vitalism, 2, 15, 24, 36 Voit, Carl, 35, 37 Volterra, Vito, 250 W Wallace, Alfred Russel, 219–224 Water, biodiversity and, 213, 214 Watson, James, 25 Wentworth, D’Arcy, 4 West, Geoffrey, 87–89, 90–92, 102–103, 107–108, 111–112, 115–116, 155–156, 236, 243, 245 West, Louis Jolyon, 67–68, 69 Willdenow, Carl Ludwig, 206–207 Wilson, E. O., 193–194 X Xylem, 112–113, 163–164 Y Yoda, Kyoji, 154, 155