Birkbeck College (University of London) School of Mathematics, Statistics and Econometrics Graduate Diploma in Financial Engineering

Applied Statistics and Econometrics Professor Ron Smith

Student Name: Bader AL-dabaan

Scaling Laws in Biology

Abstract
Do biological phenomena obey underlying universal scaling laws of life that can be mathematized so that biology can be formulated as a predictive, quantitative science ? Recent developments in Biology do indicate that this is the case. In this project we will analyse some data related with the heart rate and the lifespan as a function of the mass of animals of various scales and will nd the related regression formula. We discuss the interpretation and importance of the results.

Description

Life is certainly one of the most diverse, complex and fascinating physical phenomena in nature. The life processes cover more than 27 orders of magnitude in mass scales, from molecules of the DNA whales to sequoias. Also the corresponding metabolic rate that is required to support life across the above scales, spans over 21 orders of magnitude[1, 2, 3, 4, 5]. Despite this apparent complexity, however, observations point to the direction that, it is reasonable to conjecture that the coarse-grained behavior of living systems might obey quantiable universal laws that capture the systems essential features. For example, from the vast data available it can be found that the lifetime of a species increases as mass to the one quarter and heart rate decreases as mass to the one quarter. That leads to the fascinating conclusion that the total number of heart beats (in lifetime), is the same across all species (e.g. mammals). These scaling Laws are believed to be a manifestation of the universal underlying dynamics and geometry of the physical world. From the theoretical viewpoint, one can construct idealized biological systems whose average properties are calculable. Such ideal constructs would provide a zeroth-order point of departure for quantitatively understanding real biological systems, which can be viewed as manifesting higher-order corrections due to local environmental conditions or historical evolutionary divergence.

1.1

Recent Trends in Biology

In more detail, in biology, the observed scaling is typically a simple power law: Y = Y0 M b , where Y is some observable, Y0 a constant, and M the mass of the organism. Of even greater signicance is the fact that, the exponent b almost invariably approximates a simple multiple of 1/4 [1]. Among the many fundamental variables that obey such scaling laws are metabolic rate, life span, growth rate, heart rate, DNA nucleotide substitution rate, lengths of aortas and genomes, tree height, mass of cerebral grey matter, density of mitochondria, and concentration of RNA. In this project we intend to analyse data related with the heart rate and the lifespan as a function of the mass of animals of various scales and nd the related regression formula. We also intend to discuss the interpretation and importance of the results.

Animal Longevity and Scale

We are interested in the analysis of the eect of scale changes for animals which are similar in shape and only dier in scale. As the scale L of an animal increases the body weight and volume increase with the and hence L M 1/3 . In the following we are going to consider the eect of the change of scale through its dependence on mass. cube of scale: i.e. V L3 . Now the mass M of an animal is proportional to its volume

Scope of the Project

The main aim of this project it to determine, by using the method of least squares, estimates for the exponents (b) of the aforamentioned scaling laws. We present three sets of data: In the rst section we are investigating the relation between the average number of heart beats of various animals with their mass. In the second the average lifetime of another set of animals again as a function of their mass and lastly we have the the data related to the average lifetime heart beats and the mass of the animals. In all three cases we perform the least square method for the determination of the coecient b. We also evaluate the standard error of the regression, the determination coecient R2 and we construct the 95% condence interval for the parameter of interest b where useful. In the section corresponding to the set of data, we perform a Hypothesis Test. The general outline of our approach is the following: Given that our regressions are expected to be of the form: Y = Y0 M b (1)

Where Y is the our depedent variable, (e.g. heartbeat, longevity etc. ) of the animal, M the indepedent variable (e.g. mass, scale etc.) and Y0 , b the tting parameters. Given the non-linear nature of the regression, in order to use the method of least squares we linearize the previous by taking the Logarithm: LogY = LogY0 + bLogM (2)

now setting: y = LogL, x = LogM, a = LogL0 our regression has the form: y = a + bx (3)

in this form, we can apply the method of least squares in order to estimate the parameters a and b from the formulae (note that a, b are estimates however we drop the hat notation): a = y b x and b= The Standard Error of the regression is s2 = and the determination coecient: R2 = 1 Where u2 i (yi y )2 (7) u2 i T 2 (6) (xi x)(yi y ) 2 (xi x) (5) (4)

u2 are the residual sum of squares. Note that all coecients are estimated, i

however, for convenience we do not use the hat notation.

Heart Rates and Size

In this section we apply the method of least squares in order to determine the scaling law that relates the heart reate and the weight with the previously described approach. The data that we are going to use are given in the table below:

Animal Human Cat Small dog Medium dog Large dogs Hamster Chick Chicken Monkey Horse Cow Pig Rabbit Elephant Girae Large whales

Average Heart Rate (beats per minute) 60 150 100 90 75 450 400 275 192 44 65 70 205 30 65 20

Weight (grams) 90000 2000 2000 5000 8000 60 50 1500 5000 1200000 800000 150000 1000 5000000 900000 120000000

From the above data we can calculate the following quantities that are needed for the estimation of the parameters of the regression. For the mean of ys we have (for N = 16) y= for the mean of xs we nd, x= for the x-sum of squares we nd
N N i N i

yi

= 1.99

(8)

xi

= 4.39

(9)

x2 = 357.42 i
i

(10)

and the corresponding sum for the ys

N 2 yi = 65.84 i N

(11)

yi xi = 130.31
i

(12)

that gives b = 0.202 4 (13)

with S.E.(b) = 0.03. The intercept can be found to be: a = 2.79 (14)

4.1

Condence Interval

Given the above, we can nd the 95% ( = 5%, N = 16) condence interval for the estimate of b: b t/2,N 2 S.E.(b) (15)

value 1/4 which is expected by the idealized theoretical models (to the rst approximation) [1]. The ndings can be considered compatible with the expectations given the limitations of our sample population data.

This can be found to be b E (0.143, 0.260). Note that this interval clearly includes the

Analysis of the Regression

From the above data we have that the regression of the logarithm of Heart Rate on the logarithm of weight yields the following equation: Log(HeartRate) = 2.79 0.202Log(W eight) Or otherwise (where Heart Rate=H.R. , Weight=M), HR = AM 0.202 (17) (16)

Thus the net eect of scale (mass) on animal heart rate is negative. In more details the regression formula means that if the mass of an animal is increased by a factor of 10 then its heart rate will drop to the 0.63 of the original. In terms of scale, given that mass is proportional to the cube of the scale, we have: (HeartRate) = A(Scale)0.606 (18)

5.1

Interpretation

There have been various attempts to interpret the above. The most promising explanations are based on the fact the life is supported by chemical reactions which need oxygen to take 5

place. That brings blood ow at the center of the interest. The argument goes as following: The volume of blood ow required to feed the mass of an animal increases with the cube of scale. However, the cross sectional area of the arteries and the veins required to carry that blood ow only increases with the square of scale. There are other area-volume relationships which impose limitations on animals. The volume of the heart of an animal is proportional to the cube of scale. The volume of the blood to be moved is also proportional to the cube of scale. Putting together the previous arguments, a more detailed analysis can show that the ow velocity is approximately proportional to scale. Therefore the time required to evacuate the hearts volume is proportional to scale. This means that the heartbeat rate is approximately inversely proportional to scale. Of course, this is a qualitative description and a more detailed calculation is needed in order to expalin the behaviour of the data, In fact this has already been done in the publication[1].

Lifespan and Scale

In this section we apply the method of least squares in order to determine the scaling law that relates the Lifespan and the scale (weight), again applying the same approach. The data that we are going to use are given in the table below: Lifespan versus Weight for Various animals: Animal Human Cow Duck Cat Guinea Pig Goat Pig Monkey Dog Turkey Frog Girae Snake Weight (grams) 90000 800000 2000 2000 100 30000 150000 5000 5000 15000 50 900000 100 Life Span (years) 70 22 10 15 5 15 25 25 15 5 3 25 10

6.1

Results

From the above table (for N = 13): We calculate the means: x = 50.28 and y = 14.71 the sum of squares: x2 = 214.39 i and the sum of the products xi yi = 61.35 using these we can calculate the estimates of the regression coecients: b = 0.191 with S.E.(b) = 0.040 nd a = 0.970 (24) (23) (22) (21) (20) (19)

6.2

Condence Interval

Given the above, we can nd the 95% ( = 5%, N = 1, N = 16) condence interval for the estimate of b: b t/2,N 2 S.E.(b) (25)

This can be found to be b E (0.112, 0.269) Note that this interval clearly includes the value +1/4 which is expected by the idealized theoretical models (to the rst approximation) [1]. The nding can be considered compatible with the expectations given the limitations of our sample population data.

6.3

Determination Coecient

The determination coecient can be found to be: R2 = 0.527 (26)

This can be interpreted as that approximately, 52.7 percent of the variation in the response variable (Lifespan) can be explained by the explanatory variable(Weight). The remaining 37.3 can be explained by unknown variables or inherent variability. The t can be characterized as adequate given the limiteddata provided.

6.4

Analysis

Thus the regression formula Log(Lif espan) = 0.970 + 0.191Log(W eight) (27)

Thus the net eect of scale on animal longevity is positive. Taking into account that weight is proportional to the cube of the linear scale of an animal the above equation in terms of scale would be (assuming Scale = (W eight)1/3 ): Log(Lif espan) = 0.970 + 0.573Log(Scale) imately 5.5% longer lifespan. Or otherise, L = L0 M 0.191 Where L is the lifespan of the animal and M its mass. (29) (28)

The regression implies that animals which are 10 percent larger in scale, will have approx-

Hypothesis Testing - Lifetime and Heartbeats

From the previous we have seen that the number of heart beats scale as : HeartBeats M 0.201 and that the Lifespan as: Lifetime Heartbeats and Animal Size Lif espan M +0.191 to be independent of the mass. Let us examine this in more details: The data that we are going to use in this section are given in the table below: 8 (31) (30)

where M is the mass of the animal. Note that the product (Heartbeats)*(Lifespan) seems

Animal Human Cat Small dog Medium dog Large dogs Hamster Chicken Monkey Horse Cow Pig Rabbit Elephant Girae Large whale

Weight (gr) 90000 2000 2000 5000 8000 60 1500 5000 1200000 800000 150000 1000 5000000 900000 120000000

Heart Rate(/min) Longevity(years) 60 70 150 15 100 10 90 15 75 17 450 3 275 15 190 15 44 40 65 22 70 25 205 9 30 70 65 20 20 80

Product 4200 2250 1000 1350 1275 1350 4125 2850 1760 1430 1750 1845 2100 1300 1600

Heartbeats(106 ) 2.21 1.18 0.53 0.71 0.67 0.71 2.17 1.50 0.93 0.75 0.92 0.97 1.1 0.68 0.84

Although the lack of dependence is clear from the above data we can perform once again the method of least squares for a conrmation in terms of regression analysis: The regression can be found to be: Log(Lif etimeHeartbeats) = 9.063 0.004Log(M) The determination coecient can be calculated: R2 = 0.0019 (33) (32)

This a particularly low value of the determination coecient is a consequence of the fact that there no variation in the response variable and thus no linear relationship (or otherwise the slope is zero) between the response variable by the explanatory variable. At this stage as an simple exercise, we would like to perform a Hypothesis Test testing the our belief that the lifetime heart beats is roughly the same across species. For this we construct the null and alternative hypothesis: H0 (NullHypothesis): The number of lifetime heartbeats of animals is independent of their size (b = 0) H1 AlternativeHypothesis : The number of lifetime heartbeats of animals is independent of their size b = 0 (two tail test) 9

In order to perform the hypothesis testing we calculate the t parameter for the coecient b: b = 0.15 (34) t= S.E.(b) From the tables we nd that the relevant critical value for t for = 0.05 and degrees of freedom = 15 2 = 13 is tcr = 2.16. Given that in the present case t < tcr , we conclude that we cannot reject the null hypothesis and that there is strong evidence against the alternative hypothesis. That is to say, that the slope coecient b is insignicant , conrming that there is no dependence of lifetime heartbeats on the scale of animal size.

Summary and Conclusions

We believe that the above, indicate a deep connection between biological systems and the physical physical laws of nature. The understanding the origins of these emergent scaling laws, will potentially uncover the origin and dynamics of living systems from molecules to the biosphere. This, along with the explanation of human consciousness, is one of the most interesting challenges of modern science. By nding the universal underlying principles that govern lifes diversity we will understand the very nature of life and its emergence from rst principles. It is clear that many other disciplines, such as medicine, agriculture, and the environment will benet from these explanations. Our results where in agreement with the general expectations, however can be be improved by enlarging and diversifying the samples populations. The present, is a eld where the statistical analysis of the data is essential in order to get some insight to the cause of the emergence of these particular scaling laws. Further directions of research include the examination of additional features of animals in various scales and derive possible new scaling laws. From the theoretical point of view one can construct models that try to explain the phenomenology from physical and chemical reasoning. There is some signicant progress in this direction[1].

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References
[1] J. H. Brown, G. B. West, eds., Scaling in Biology, Oxford U. Press, New York (2000)

[2] T. A. McMahon, J. T. Bonner, On Size and Life, Scientic American Library, New York (1983) ; [3] K. Schmidt-Nielsen, Scaling: Why Is Animal Size So Important?, Cambridge U. Press, New York (1984);

[4] R. H. Peters, The Ecological Implications of Body Size, Cambridge U. Press, New York (1983); W. A. Calder III, Size, Function and Life History, Harvard U. Press, Cambridge, MA (1984)

[5] K. J. Niklas, Plant Allometry: The Scaling of Form and Process, U. of Chicago Press, Chicago (1994);

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