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Scaling laws

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How quantities scale, with the exponent, the scope, and the falsifier for each.

68 rows.

name relation exponent scope class source falsifier
b b = ≈ 0.74 (Brody, same year: ≈ 0.73) ≈ 0.74 (Brody, same year: ≈ 0.73) ~13 spp mammals + birds, 1932 dataset OBSERVED-CONTESTED Kleiber (1932), Hilgardia 6:315–353 See the dispute row below.
b_WBE b_WBE = 3/4 = 0.75 exactly 3/4 = 0.75 exactly conditional on: space-filling fractal branching network + size-invariant terminal units + minimised dissipation MODELED West, Brown & Enquist (1997), Science 276(5309):122–126, DOI 10.1126/science.276.5309.122 Exhibit a taxon meeting all three assumptions whose measured b excludes 0.75. Drop any assumption and 3/4 does not follow.
b_WS b_WS = 0.686 ± 0.014 (95% CI); interordinal 0.65 0.686 ± 0.014 (95% CI); interordinal 0.65 571 mammal spp; BMR normalised to 36.2 °C; excl. Artiodactyla, Macropodidae, Lagomorpha, Soricidae; interspecific ≈0.69, N=619, r²=0.94 OBSERVED-CONTESTED White & Seymour (2003), PNAS 100(7):4046–4049, DOI 10.1073/pnas.0436428100; Dodds, Rothman & Weitz (2001), J. Theor. Biol. 209(1):9–27 Pre-register taxon set + temperature normalisation + regression unit, and show the CI is stable across all three. It currently is not — that is the dispute.
b_Kleiber b_Kleiber = 3/4 (measured slope reported 0.74) 3/4 (measured slope reported 0.74) BMR vs mass, 13 data points OBSERVED-CONTESTED Kleiber 1932, Hilgardia 6:315–353; n=13 per Kolokotrones et al. 2010, Nature 464:753–756 See every row below; the dispute IS the falsification record.
b_Rubner b_Rubner = 2/3 2/3 surface-law argument; respiration trials on dogs OBSERVED-CONTESTED Rubner 1883 Data rejecting 2/3 at a stated mass range.
b_Savage,binned b_Savage,binned = 0.737 0.737 mammal BMR, 0.1 log-unit bins OBSERVED-CONTESTED Savage et al. 2004, Funct Ecol 18:257–282 (95% CI 0.711–0.762, n=52) Re-fit with different binning.
b_Savage,unbinned b_Savage,unbinned = 0.712 0.712 mammal BMR, all species — CI excludes 2/3 and 3/4 OBSERVED-CONTESTED Savage et al. 2004 (95% CI 0.699–0.724, n=626) Re-fit; show binning is not what moves the verdict.
b_White&Seymour b_White&Seymour = 0.68 (interspecific); 0.65 (interordinal) — both ≠ 3/4, both = 2/3 within error 0.68 (interspecific); 0.65 (interordinal) — both ≠ 3/4, both = 2/3 within error 619 spp. → 469 after excluding Artiodactyla, Lagomorpha, Soricidae, Macropodidae; T_b-corrected to 36.2 °C, Q₁₀=3.0 OBSERVED-CONTESTED White & Seymour 2003, PNAS 100(7):4046–9 (CIs not read in this pass) A dataset with equivalent basal-condition rigour giving 3/4.
b_WS,BMR / SMR / RMRt b_WS,BMR / SMR / RMRt = 0.686±0.014 / 0.675±0.013 / 0.712±0.013 0.686±0.014 / 0.675±0.013 / 0.712±0.013 exponent depends on which rate is measured; the BMR/SMR figures are White & Seymour's own 2003 fits restated in their 2005 review — same authors, same dataset, same regression, so not a replication OBSERVED-CONTESTED (re-classed: the identical underlying result is OBSERVED-CONTESTED two rows up; one result cannot hold two classes) White & Seymour 2003, as tabulated in White & Seymour 2005, J Exp Biol 208:1611 Show the three definitions give one exponent.
b₂ (curvature) b₂ (curvature) = 0.0322 ± 0.0053 (P = 9.0×10⁻¹⁰); 0.0294 ± 0.0057 with T 0.0322 ± 0.0053 (P = 9.0×10⁻¹⁰); 0.0294 ± 0.0057 with T McNab dataset, n=636 (447 with T); unit-scale invariant OBSERVED-CONTESTED Kolokotrones et al. 2010, Nature 464:753–756, Table 1 MacKay 2011 — see contested row; reply: Deeds, Savage & Fontana 2011.
b₁ b₁ = 0.5400 ± 0.0295 (0.5371 ± 0.0305 with T) 0.5400 ± 0.0295 (0.5371 ± 0.0305 with T) artifact of M₀ = 1 g; not interpretable alone INADMISSIBLE as 'the exponent' Kolokotrones et al. 2010; objection: MacKay 2011, J Theor Biol 280(1):194–6; reply: Deeds, Savage & Fontana 2011, 280:197–8 — concedes the artifact, calls it irrelevant to curvature; the class stands either way Derive: under M'=kM, b₁' = b₁ − 2b₂log k.
local slope local slope = 0.57 → 0.87 (rises with mass) 0.57 → 0.87 (rises with mass) ~3.6 g to ~460 kg; = b₁ + 2b₂log₁₀M MODELED Computed in-chapter from Kolokotrones Table 1; matches their stated range Arithmetic error.
M(slope=1) M(slope=1) = ~7.4 × 10⁷ g ≈ 74 t ~7.4 × 10⁷ g ≈ 74 t proposed upper bound on animal size — EXTRAPOLATION: ~2.2 decades beyond the fitted data, which end at ~460 kg (local slope 0.87). Kolokotrones et al.'s own two hedges: the unbounded slope rise 'may be due to the paucity of data for large animals', and the size-limit reading holds only 'If this is correct' MODELED (extrapolated — rule 1 applies to this row) Computed in-chapter; Kolokotrones et al. state ~10⁸ g (100 t) A larger animal; or the slope not reaching 1.
k_marsupial k_marsupial = 0.75 ± 0.01, R² = 0.990 0.75 ± 0.01, R² = 0.990 70 marsupials (McNab 2008), excluding Tarsipes rostratus + Lasiorhinus latifrons OBSERVED-CONTESTED MacKay 2011 Re-fit with the 2 species retained.
WBE assumptions WBE assumptions = space-filling fractal network; size-invariant terminal unit; energy minimisation space-filling fractal network; size-invariant terminal unit; energy minimisation derivation of 3/4 MODELED West, Brown & Enquist 1997, Science 276:122–126 The assumptions are the fence — see rows below.
WBE finite-size form WBE finite-size form = M = c₀B + c₁B^(4/3), both c > 0 → concave; data are convex M = c₀B + c₁B^(4/3), both c > 0 → concave; data are convex wrong sign of curvature MODELED (refuted on this point) Kolokotrones et al. 2010 Show c₁ < 0 follows from WBE's own minimisation.
b_heart b_heart = −1/4 −1/4 resting mammals; L&H state no n and no mass range for the resting-HR claim — their 34-species / 7 g–500 kg series is maximal heart rate (−0.15) and VO₂max, not this row OBSERVED-REPLICATED Lindstedt & Hoppeler 2023, J Exp Biol 226(24):jeb245766 — 'resting heart rate scales as M–1/4' Modern re-fit with CI outside −0.30 to −0.20.
b_lifespan,mammal b_lifespan,mammal = 0.153 (t_max = 4.88·M^0.153 yr, M in g), R²=0.66 0.153 (t_max = 4.88·M^0.153 yr, M in g), R²=0.66 856 mammals, cetaceans excluded — not 1/4 OBSERVED-REPLICATED de Magalhães, Costa & Church 2007, J Gerontol A 62(2) Re-fit giving CI containing 0.25.
b_lifespan,bird b_lifespan,bird = 0.218 (t_max = 5.22·M^0.218 yr), R²=0.70 0.218 (t_max = 5.22·M^0.218 yr), R²=0.70 518 birds OBSERVED-REPLICATED de Magalhães et al. 2007 ('body mass explained 70% of the variation in tmax') As above.
b_VO₂max b_VO₂max = 0.872 0.872 34 eutherian species, 7 g – 500 kg — vs basal ~0.70 OBSERVED-REPLICATED Lindstedt & Hoppeler 2023 Show basal and max share an exponent.
beats/lifetime scaling beats/lifetime scaling = ∝ M^(−0.097) → ~4.8× decline over 7 decades of mass ∝ M^(−0.097) → ~4.8× decline over 7 decades of mass composition of −0.25 and +0.153 MODELED (computed in-chapter) Computed from Lindstedt & Hoppeler 2023 + de Magalhães et al. 2007 Measure beats/lifetime vs mass directly in one dataset — this composition mixes sources (M22).
σ ∝ L σ ∝ L = stress grows linearly with size at constant shape stress grows linearly with size at constant shape F/A ∝ L³/L² OBSERVED-REPLICATED (geometry) Galileo 1638, Two New Sciences Geometric error.
elastic similarity elastic similarity = L ∝ D^(2/3); → D ∝ M^(3/8), L ∝ M^(1/4), S ∝ M^(5/8) L ∝ D^(2/3); → D ∝ M^(3/8), L ∝ M^(1/4), S ∝ M^(5/8) McMahon's model MODELED McMahon 1973, Science 179:1201–4; cascade computed in-chapter (reproduces his M^(5/8)) See next row — largely refuted empirically.
bone scaling (measured) bone scaling (measured) = length ∝ M^0.31; diameter ∝ M^0.35 length ∝ M^0.31; diameter ∝ M^0.35 32 mammal spp. (secondary sources say 37 — unresolved here), 0.020–3500 kg — close to geometric similarity, not elastic (0.25 / 0.375) OBSERVED-REPLICATED (primary not read in this pass; exponents and n via secondary sources) Alexander et al. 1979, J Zool 189:305–314 Read the primary; a species count or exponent outside the stated values moves this row. The NEGATIVE verdict rests on the direction (geometric, not elastic), which is corroborated independently, and survives either count.
b_colony,metabolic b_colony,metabolic = 0.81, 95% CI 0.55–1.08 0.81, 95% CI 0.55–1.08 12 colonies + 391 unitary insects; CI excludes nothing OBSERVED-CONTESTED Hou et al. 2010, PNAS 107(8):3634–8 More colonies; a CI that excludes an alternative.
b_colony,production b_colony,production = 0.74, 95% CI 0.71–0.76 (r²=0.99, combined) 0.74, 95% CI 0.71–0.76 (r²=0.99, combined) colonies + unitary organisms — the tight row OBSERVED-REPLICATED Hou et al. 2010 Independent re-fit.
b_city,superlinear b_city,superlinear = cluster 1.07–1.34, not a single value cluster 1.07–1.34, not a single value patents 1.27 [1.25–1.29]; R&D empl. 1.34 [1.29–1.39]; GDP 1.15 [1.06–1.23]; wages 1.12 [1.09–1.13]; AIDS 1.23 [1.18–1.29] OBSERVED-CONTESTED Bettencourt et al. 2007, PNAS 104(17):7301–6, Table 1 Leitão et al. 2016 — see next row.
b_city,sublinear b_city,sublinear = gasoline stations 0.77 [0.74–0.81]; road surface 0.83 [0.74–0.92] (n=29); cables 0.87 [0.82–0.92] gasoline stations 0.77 [0.74–0.81]; road surface 0.83 [0.74–0.92] (n=29); cables 0.87 [0.82–0.92] Germany/USA 2001–02 OBSERVED-CONTESTED Bettencourt et al. 2007 As above.
urban β ≠ 1 urban β ≠ 1 = model-dependent model-dependent 5 models × 15 datasets; depends on fluctuations, their model, and heavy-tailed city sizes OBSERVED-CONTESTED Leitão et al. 2016, R Soc Open Sci 3:150649 (arXiv:1604.02872) A fluctuation model class under which the verdict is stable.
Bergmann conformity Bergmann conformity = 65–71% (mammals); 72–76% (birds) 65–71% (mammals); 72–76% (birds) 149 mammals, 94 birds OBSERVED-CONTESTED Meiri & Dayan 2003, J Biogeogr 30:331–351 — percentages via Teplitsky & Millien 2014; primary not read (M22) Read the primary; a value outside these ranges.
exponent spread exponent spread = mean 0.738±0.018 but 51% of exponents outside 0.7–0.8; range <0.5 to >1.0 mean 0.738±0.018 but 51% of exponents outside 0.7–0.8; range <0.5 to >1.0 146 relations (Peters 1983), 72% vertebrate OBSERVED-REPLICATED Glazier 2005, Biol Rev 80:611–662 Recount the distribution.
intraspecific spread intraspecific spread = 0.3 to 1.8; mean 0.724, mode 0.667 0.3 to 1.8; mean 0.724, mode 0.667 220 species (Withers 1992) OBSERVED-REPLICATED Glazier 2005 Recount.
MLBH bounds MLBH bounds = 2/3 (surface-area limits) to 1 (mass/volume power limits) 2/3 (surface-area limits) to 1 (mass/volume power limits) metabolic-level boundaries hypothesis HYPOTHESIZED Glazier 2005, 2010, Biol Rev 85:111–138 An exponent stably outside [2/3, 1] with a demonstrated mechanism.
Metabolic/rate exponent Metabolic/rate exponent = 2/3 vs 3/4 — disputed 2/3 vs 3/4 — disputed mammals; lineage-dependent OBSERVED-CONTESTED 3/4: Kleiber-family. 2/3: White & Seymour (2003) PNAS 100:4046–4049 (619 spp.); Dodds, Rothman & Weitz (2001) JTB; lineage-varying: Capellini, Venditti & Barton (2010) Ecology 91(9) A phylogenetically-controlled dataset where all lineages converge on one exponent would settle it.
t(1 m) t(1 m) = ~6.2 × 10⁹ (≈195 yr) ~6.2 × 10⁹ (≈195 yr) 3D diffusion, D = 27 µm²/s, t = L²/6D MODELED Computed in-chapter from BNID 101997 + <r²>=6Dt Arithmetic error, or a demonstration of 1 m protein transport by diffusion alone.
S/V S/V = 6 vs 0.3 6 vs 0.3 sphere 3/R; R = 0.5 µm vs 10 µm MODELED Computed in-chapter (geometry) Geometric error.
δc/c scaling δc/c scaling = ∝ (D·a·c·T)^(−1/2) ∝ (D·a·c·T)^(−1/2) diffusion-limited chemoreception OBSERVED-REPLICATED (as a scaling) Berg & Purcell 1977, Biophys J 20:193–219 A sensor beating the −1/2 exponent.
δc/c prefactor δc/c prefactor = disputed disputed B–P vs Bialek–Setayeshgar vs Kaizu: B–S term missing 1/(2(1−n̄)) OBSERVED-CONTESTED / MODELED Bialek & Setayeshgar 2005, PNAS 102(29):10040–5; Kaizu et al. 2014, Biophys J 106(4):976–85 A treatment retaining receptor–ligand correlations that settles the constant.
b_prokaryote b_prokaryote = 1.7 active / 2.0 inactive 1.7 active / 2.0 inactive metabolic rate vs body mass; n = 44 / 121 OBSERVED-REPLICATED DeLong et al. 2010, PNAS 107(29):12941–5, DOI 10.1073/pnas.1007783107 Refit with independent data; CI covering 0.75.
b_protist b_protist = 1.0 active / 1.1 inactive 1.0 active / 1.1 inactive n = 51 / 52 OBSERVED-REPLICATED DeLong et al. 2010 As above.
b_metazoan b_metazoan = 0.76 active / 0.79 inactive 0.76 active / 0.79 inactive n = 71 / 15 OBSERVED-REPLICATED DeLong et al. 2010 As above.
Kleiber universality Kleiber universality = refuted as universal refuted as universal 3/4 does not apply across prokaryote/protist/metazoan OBSERVED-REPLICATED DeLong et al. 2010 (their explicit conclusion) A dataset in which one exponent fits all three groups.
Metabolic curvature Metabolic curvature = convex on log-log; quadratic in log-mass required; not a pure power law convex on log-log; quadratic in log-mass required; not a pure power law mammals, temperature-corrected OBSERVED-CONTESTED Kolokotrones et al. 2010, Nature 464:753–6, DOI 10.1038/nature08920; contested by MacKay 2011, J Theor Biol 280(1):194–196; replied to by Deeds, Savage & Fontana 2011, J Theor Biol 280(1):197–198 — the exchange is live on both sides Quadratic coefficient CI covering zero on independent data.
Exponent-by-subset Exponent-by-subset = small-dominated → ~2/3; large-dominated → ~3/4 small-dominated → ~2/3; large-dominated → ~3/4 mammals; artefact of fitting a line to a curve OBSERVED-CONTESTED (with the row above) Kolokotrones et al. 2010 Both subsets return the same slope.
α ∝ f² α ∝ f² = exponent 2 exponent 2 classical (Stokes–Kirchhoff) term only MODELED classical acoustics Not the observed exponent below 10 kHz — see next rows.
Rayleigh Rayleigh = ∝ 1/λ⁴ ∝ 1/λ⁴ scatterers ≪ λ OBSERVED-REPLICATED Strutt [Rayleigh] 1871, Phil. Mag. 41:107–120, 274–279 A small-particle scattering exponent ≠ 4.
c ∝ √T c ∝ √T = exponent ½ exponent ½ ideal gas; independent of pressure MODELED Computed in-chapter from √(γRT/M) Demonstrate a pressure dependence of c at fixed T.
W/S ∝ m^(1/3); v ∝ m^(1/6) W/S ∝ m^(1/3); v ∝ m^(1/6) = exponents ⅓, ⅙ exponents ⅓, ⅙ isometric scaling of a flyer MODELED Computed in-chapter Show non-isometric wing-area scaling that breaks it.
P_req ∝ m^(7/6) P_req ∝ m^(7/6) = exponent 7/6 exponent 7/6 induced power, isometric MODELED Computed in-chapter; = Pennycuick's 7/6 law Arithmetic; or measured power scaling ≠ 7/6.
P_avail ∝ m^(2/3) P_avail ∝ m^(2/3) = exponent 2/3 exponent 2/3 muscle mass ∝ m, wingbeat frequency falls with size MODELED Computed in-chapter; the second curve in the crossing Measured power-available scaling ≠ 2/3.
max flapping mass (theory) max flapping mass (theory) = ~12 ~12 aerobically powered continuous flapping flight; m^(7/6) vs m^(2/3) crossing MODELED (NOT-SOURCED — via secondary reports, not the primary) Pennycuick's aerodynamic theory; secondary reports also give 'largest extant flying species ≈12–14 kg' (Pennycuick 1989) Fetch the primary; or a measured power-available exponent ≠ 2/3.
ceiling vs census ceiling vs census = exceedance, not agreement exceedance, not agreement heaviest bustards sit above the ~12 kg theoretical ceiling OBSERVED-CONTESTED this chapter; the two rows above, each in its own scope Show a >16 kg bird sustaining continuous aerobic flapping → refutes the ceiling. Show bustard flight is burst-only → the ceiling's scope condition holds and the exceedance is not one.
α (MLR) α (MLR) = 2.028 / 4.572 / 5.743 / 4.329 / 3.967 / 2.865 2.028 / 4.572 / 5.743 / 4.329 / 3.967 / 2.865 six pieces over 0.179–31 M☉; 509 stars, detached eclipsing binaries OBSERVED-REPLICATED Eker et al. 2018, MNRAS 479(4):5491–5511, doi:10.1093/mnras/sty1834 (Table 4) An independent DEB sample with a single exponent, or different break points.
α = 3.5 α = 3.5 = not found in any piece not found in any piece the textbook value SUPERSEDED contradicted by Eker et al. 2018 Show a mass range where 3.5 is the calibrated fit.
t_MS(30 M☉) t_MS(30 M☉) = 2 × 10⁶ (α=3.5) vs 1.8 × 10⁷ (α=2.865); robust range 10⁶–10⁷ 2 × 10⁶ (α=3.5) vs 1.8 × 10⁷ (α=2.865); robust range 10⁶–10⁷ t ∝ M^(1−α), anchored on the Sun MODELED Computed in-chapter from Eker et al. 2018 + t ∝ M/L A detailed evolutionary track for 30 M☉ outside 10⁶–10⁷ yr.
b_geom b_geom = 1.0 1.0 predicted length-vs-circumference exponent, geometric similarity MODELED Kilbourne & Makovicky 2010, J Anat, Table 8 Derivation error.
b_elastic b_elastic = 0.67 0.67 elastic similarity (L ∝ D^(2/3)); attributed to McMahon 1975a MODELED Kilbourne & Makovicky 2010, Table 8 Derivation error.
b_stress b_stress = 0.5 0.5 static stress similarity MODELED Kilbourne & Makovicky 2010, Table 8 Derivation error.
b_fem,Trex b_fem,Trex = 0.5341 — 95% CI 0.04159–0.9718 0.5341 — 95% CI 0.04159–0.9718 T. rex, femur, ontogenetic, RMA log L vs log C — CI contains static-stress (0.5), elastic (0.67) and nears geometric (1.0): discriminates among NONE; carries no contrast with any other taxon OBSERVED-SINGLE (one growth series, one study; CI spans the model space — downgraded from OBSERVED-REPLICATED) Kilbourne & Makovicky 2010, J Anat, Table 3 Re-measure the growth series; RMA slope or CI outside the published 0.04159–0.9718.
b_fem,Allo b_fem,Allo = 0.82 0.82 Allosaurus fragilis, femur, ontogenetic OBSERVED-REPLICATED Kilbourne & Makovicky 2010 As above.
b_fem,sauropodomorph b_fem,sauropodomorph = ~1.0 (Massospondylus 0.81) ~1.0 (Massospondylus 0.81) sauropodomorphs, femur, ontogenetic OBSERVED-REPLICATED Kilbourne & Makovicky 2010 As above.
b_fem,hadrosaur b_fem,hadrosaur = 1.05 (Maiasaura); 1.094 (Hypacrosaurus, 95% CI 1.072–1.113) 1.05 (Maiasaura); 1.094 (Hypacrosaurus, 95% CI 1.072–1.113) hadrosaurids, femur, ontogenetic — tight CI excludes every standard model from below; this is the section's real signal, and it stands alone without the T. rex row OBSERVED-REPLICATED Kilbourne & Makovicky 2010, Table 3 Re-measure; slope or CI overlapping 1.0.
b_fem,interspecific b_fem,interspecific = 0.83 (tibia 0.78; MT III 0.80) 0.83 (tibia 0.78; MT III 0.80) interspecific, non-avian dinosaurs — do not conflate with ontogenetic OBSERVED-REPLICATED Carrano, as reported by Kilbourne & Makovicky 2010 Fetch Carrano primary; femoral exponent outside ~0.83 ± 0.05.
EE_odontocete EE_odontocete = decreases with body mass decreases with body mass energy captured ÷ energy expended OBSERVED-REPLICATED Goldbogen et al. 2019, Science 366:1367–1372 Tag data showing EE rising with size in odontocetes.
EE_rorqual EE_rorqual = increases with body mass increases with body mass lunge filter feeders on krill OBSERVED-REPLICATED Goldbogen et al. 2019, Science As above, inverted.
v ∝ d v ∝ d = linear linear conduction velocity vs outer diameter, myelinated OBSERVED-REPLICATED Hursh 1939, Am J Physiol 127(1):131–139; Gasser & Grundfest 1939 A myelinated preparation with non-linear v(d).
W ∝ G^(4/3) W ∝ G^(4/3) = 4/3 (less a small cortical-thickness correction) 4/3 (less a small cortical-thickness correction) white vs grey matter, mammals, several orders of magnitude OBSERVED-REPLICATED Zhang & Sejnowski 2000, PNAS 97:5621–5626 A mammalian dataset fitting a materially different exponent.
V_wire ∝ R⁶ V_wire ∝ R⁶ = 6 6 crude model: wiring volume to hold τ fixed as R grows; assumes fixed fraction of long-range neurons, linear v(d), spherical brain MODELED Computed in-chapter; cf. Zhang & Sejnowski's more careful 4/3 derivation The assumptions — chiefly fixed fraction of long-range neurons, which modular brains violate by design.

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