Allometric exponents

support a 3/4 power scaling

law

Catherine C. Farrell
Nicholas J. Gotelli
Department of Biology
University of Vermont
Burlington, VT 05405
Gotelli lab, May 2005
 Allometric Scaling
• What is the relationship metabolic rate (Y)
and body mass (M)?
 Allometric Scaling
• What is the relationship metabolic rate (Y)
and body mass (M)?
• Mass units: grams, kilograms
• Metabolic units: calories, joules, O2
consumption, CO2 production
 Allometric Scaling
• What is the relationship metabolic rate (Y)
and body mass (M)?
• Usually follows a power function:
• Y = CMb
 Allometric Scaling
• What is the relationship metabolic rate (Y)
and body mass (M)?
• Usually follows a power function:
• Y = CMb
• C = constant
• b = allometric scaling coefficient
 Allometric Scaling: Background
• Allometric scaling equations relate basal
metabolic rate (Y) and body mass (M) by
an allometric exponent (b)

Y = YoMb Log Y = Log Yo + b log M

12 4.5
4
10
3.5
8 3
2.5
Y 6 Log (Y)
2
4 1.5

2 1
0.5
0
0
0 20 40 60 80 100 120 0 0.5 1 1.5 2 2.5
M Log (M)
 Allometric Scaling: Background
• Allometric scaling equations relate basal
metabolic rate (Y) and body mass (M) by
an allometric exponent (b)

Y = YoMb Log Y = Log Yo + b log M

12 4.5
4
10
3.5
8 3
2.5
Y 6 Log (Y)
2
4 1.5

2 1
0.5
0 b is the slope of
0
0 20 40 60
M
80 100
the log-log plot!0
120 0.5 1 1.5 2 2.5
Log (M)
 Allometric Scaling
• What is the expected value of b?

4.5
4
3.5
3
2.5
Log (Y)
2
1.5
1
0.5
?? 0
0 0.5 1 1.5 2 2.5
Log (M)
Hollywood Studies Allometry

Godzilla (1954)
A scaled-up dinosaur
Hollywood Studies Allometry

The Incredible Shrinking Man (1953)

A scaled-down human
 Miss Allometry

Raquel Welch
Movies spanning > 15 orders of
magnitude of body mass!
1 Million B.C. (1970)
Fantastic Voyage (1964)
Hollywood (Finally)
Learns Some Biology

Alien (1979) Antz (1998)

Hollywood’s Allometric Hypothesis:
4.5
4
3.5
3
2.5
Log (Y)
2
1.5
1
0.5
b = 1.0 0
0 0.5 1 1.5 2 2.5
Log (M)
Surface/Volume Hypothesis
4.5
4
3.5
3
2.5
Log (Y)
2
1.5
1
0.5
b = 2/3 0
0 0.5 1 1.5 2 2.5
Log (M)

Surface area  length2 Volume  length3

Surface/Volume Hypothesis
4.5
4
3.5
3
2.5
Log (Y)
2
1.5
1
Microsoft Design
0.5
b = 2/3 0
Flaw!
0 0.5 1 1.5 2 2.5
Log (M)

Surface area  length2 Volume  length3

 New allometric theory of the 1990s

• Theoretical models of universal

quarter-power scaling relationships
– Predict b = 3/4
– Efficient space-filling energy transport
(West et al. 1997)
– Fractal dimensions (West et al. 1999)
– Metabolic Theory of Ecology (Brown
2004)
 Theoretical Predictions
• b = 3/4
– Maximize internal exchange efficiency
– Space-filling fractal distribution networks (West et al.
1997, 1999)

• b = 2/3
– Exterior exchange geometric constraints
– Surface area (length2): volume (length3)
 Research Questions
Meta-analysis of published exponents

1. Is the calculated allometric exponent (b)

correlated with features of the sample?
2. Mean and confidence interval for
published values?
3. Likelihood that b = 3/4 vs. 2/3?
4. Why are estimates often < 3/4?
Allometric Species Taxon Source
exponent in sample
0.71 391 mammals (Heusner 1991)
0.713 321 mammals (McNab 1988)
0.69 487 mammals (Lovegrove
2000)
0.737 626 mammals (Savage et al.
2004)
0.74 10 mixed (Kleiber 1932)
0.76 228 mammals (West et al.
2002)
0.724 35 passerine (Lasiewski and
birds Dawson 1967)
 Research Questions
1. Is the calculated allometric exponent (b)
correlated with features of the sample?
2. Calculate mean & confidence interval for
published values?
3. Likelihood that b = 3/4 vs. 2/3
4. Why are estimates often < 3/4?
 Question 1
• Can variation in published allometric
exponents be attributed to variation in
– sample size
– average body size
– range of body sizes measured
Allometric exponent as a function
of number of species in sample
P = 0.6491
0.90
Mammal
Allometric Exponent

0.85
s
0.80
Other
0.75

0.70

0.65

0.60
0 100 300 500 700

Number of species in sample

Allometric exponent as a function
of midpoint of mass
P = 0.5781
Weighted by sample size P = 0.565

0.90 Mammals
Allometric Exponent

0.85 Other
0.80

0.75

0.70

0.65

0.60
0 500 1000 1500 2000

Midpoint of mass
Allometric exponent as a function
of log(difference in mass)
P = 0.5792
Weighted by sample size: P = .649
0.90
Mammals
0.85
Allometric Exponent

0.80 Other
0.75

0.70

0.65

0.60
0 1 2 3 4 5 6

Log(difference in mass)
 Non-independence in Published
Allometric Exponents
• phylogenetic non-independence
– species within a study exhibit varying levels of
phylogenetic relatedness
Bokma 2004, White and Seymour 2003

• data on the same species are sometimes

used in multiple studies
 Independent Contrast Analysis
• Paired studies analyzing related taxa (Harvey and Pagel 1991)
– e.g., marsupials and other mammals
• Each study was included in only one pair
• No correlation (P > 0.05) between difference in the
allometric exponent and
– difference in sample size,
– midpoint of mass
– range of mass
 Question 1: Conclusions
• Allometric exponent was not correlated with
– sample size
– midpoint of mass
– range of body size
• Reported values not statistical artifacts
 Research Questions
1. Is the calculated allometric exponent (b)
correlated with features of the sample?
2. Calculate mean & confidence interval for
published values?
3. Likelihood that b = 3/4 vs. 2/3
4. Why are estimates often < 3/4?
 Question 2: What is the best
estimate of the allometric
exponent?

Mammals Birds Reptiles

 0.90

0.85
Allometric Exponent

0.80

0.75 b = 3/4

0.70
b = 2/3
0.65

0.60
Mammals Birds Reptiles
 0.90

0.85
Allometric Exponent

0.80

0.75 b = 3/4

0.70
b = 2/3
0.65

0.60
Mammals Birds Reptiles
 0.90

0.85
Allometric Exponent

0.80

0.75 b = 3/4

0.70
b = 2/3
0.65

0.60
Mammals Birds Reptiles
 Question 2: Conclusions
Mammals and Birds Reptiles
Results suggest the true Variation is due to small
exponent is between 2/3 sample sizes and
and 3/4 variability in
experimental conditions
 Research Questions
1. Is the calculated allometric exponent (b)
correlated with features of the sample?
2. Calculate mean & confidence interval for
published values?
3. Likelihood that b = 3/4 vs. 2/3?
4. Why are estimates often < 3/4?
Question 3: Likelihood Ratio

b = 3/4 : b = 2/3

All species 16 074

Mammals 105

Birds 7.08

Reptiles 2.20
 Research Questions
1. Is the calculated allometric exponent (b)
correlated with features of the sample?
2. Calculate mean & confidence interval for
published values?
3. Likelihood that b = 3/4 vs. 2/3?
4. Why are estimates often < 3/4?
 Question 4: estimates often < 3/4?
0.90

0.85
Allometric Exponent

0.80

0.75 b = 3/4

0.70
b = 2/3
0.65

0.60
Mammals Birds Reptiles
 Linear Regression
• Most published exponents based on linear regression
• Assumption: x variable is measured without error
• Measurement error in x may bias slope estimates
 Measurement Error

• Limits measurement of true species mean mass

• Includes seasonal variation
• Systematic variation
• “Classic” measurement errors
 Simulation: Motivation
e.g. y = 2xtrue

200 200

150 Slope = 2.0 150

100 100

50 50
Slope = 1.8
0 0
0 20 40 60 80 100 0 20 40 60 80 100 120
True measurement Error in measurement
 Simulation: Assumptions
Assumed model Y = met. Rate
Yi = mi 0.75 m = mass
Add variation in measurement X = error term (can be
of mass positive or negative)
Yi = (mi + Xi)b b = exponent
Simulate error in measurement K = % measurement error
Xi = KmiZ Z = a random number
Z ~ N(0,1)
 0.76
Allometric Exponent 0.75
0.74
0.73
0.72
0.71
0.70
0.05 0.10 0.15 0.20
Proportion Measurement Error

Circles: mean of 100 trials

Triangles: estimated parametric confidence intervals
 Question 4: Conclusions

• Biases slope estimates down

• Never biases slope estimates up
• Parsimonious explanation for discrepancy
between observed and predicted allometric
exponents for homeotherms.
 Slope Estimates Revisited

• Other methods than least-squares can be

used to fit slopes to regression data
• “Model II Regression” does not assume
that error is only in the y variable
• Equivalent to fitting principal components
 Ordinary Least-Squares Regression
• Most published exponents based on OLS
• Assumption: x variable is measured without error
• Fitted slope minimizes vertical residual deviations
from line
 Reduced Major Axis Regression
• Minimizes perpendicular distance of points to line
• Does not assume all error is contained in y variable
• “Splits the difference” between x and y errors
 Reduced Major Axis Regression
• Slope of Major Axis Regression
is always > slope of OLS Regressions
• Major Axis Regression slope = b / r2

increasing b
 Re-analysis of Data
• Adjusted slope for n = 5 mammal data sets
 Conclusions

• Measured allometric exponents not

correlated with features of sample
• Published exponents cluster tightly for
homeotherms
– values slightly lower than the
predicted b = 3/4.
• Published exponents highly variable for
poikilotherm studies
 Conclusions

• Body mass measurement error always

biases least-squares slope estimates
downward
• Observed allometric exponents closer to
3/4 than 2/3
 Acknowledgements

Gordon Research Conference Committee

Metabolic Basis of Ecology
Bates College
July 4-9, 2004