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The Journal of Experimental Biology 202, 24952503 (1999) Printed in Great Britain The Company of Biologists Limited 1999

9 JEB1959



JOHN D. CURREY* Department of Biology, University of York, York YO10 5YW, UK

Accepted 7 June; published on WWW 25 August 1999 Summary The bending strength of a wide variety of bony types is material allows a greater bending moment to be exerted shown to be nearly linearly proportional to Youngs after the yield point has been reached, thereby increasing modulus of elasticity/100. A somewhat closer and more the strength as calculated from beam formulae. (2) Loading satisfactory t is obtained if account is taken of the in bending results in a much smaller proportion of the variation of yield strain with Youngs modulus. This volume of the specimens being raised to high stresses than nding strongly suggests that bending strength is is the case in tension, and this reduces the likelihood of a determined by the yield strain. The yield stress in tension, weak part of the specimen being loaded to failure. which might be expected to predict the bending strength, underestimates the true bending strength by Key words: bone, bending strength, Youngs modulus, Weibull approximately 40 %. This may be explained by two effect. phenomena. (1) The post-yield deformation of the bone

Introduction There have been two general approaches to explaining the mechanical properties of bone. One approach is to explain the mechanical properties by reference to the microscopic and ne structure of the bone. Katz (1971) made a pioneering attempt to explain elastic properties in this way. Examples of later attempts are those by Wagner and Weiner (1992) and Sasaki et al. (1991), who sought to explain some elastic properties by considering the effect of the mineral crystal size on the properties. However, explanations of failure, as opposed to elastic, behaviour using microscopic and ne structural information have effectively all been qualitative. The other approach is to relate statistically the mechanical behaviour to other features of the bone, such as porosity, the general orientation of the bones structure or the mineral content. This is phenomenological: one can explain mechanical properties in terms of other variables and use such relationships for prediction. Keller (1994) related various compressive properties to mineral content and porosity. Martin and Ishida (1989) examined the relationship between tensile strength and mineral content, porosity, and the predominant orientation of the bone. Similarly, Currey (1987, 1990) used a large data set to examine the physical and chemical properties of compact bone, in particular its mineral content and porosity, and their effect on bones mechanical properties. The properties mainly examined were Youngs modulus of elasticity and those associated with tensile failure. In the present study, I discuss results, from an enlarged data set, for bending strength.

Materials and methods The data set The total data set consists of records from 951 specimens from 67 compact mineralised tissues from 32 species. Estimates of porosity were made for 370 of these specimens. The bending tests reported here were carried out on 351 specimens from 52 bones from 23 species (Table 1). The 86 bending specimens for which estimates of porosity were made came from 35 bones from 18 species. Not all the specimens are of bone sensu stricto, there are also some results from elephants tusk and sarus crane ossied tendon. However, all tissues will be referred to as bone. Mechanical testing Both bending and tensile specimens were prepared. The general method of preparation and testing was as described by Currey (1988). It involved rough preparation with a bandsaw, and smoothing with increasingly ne grades of carborundum paper. Tensile specimens were shaped into a dumbbell shape by a milling head guided by a pre-machined pattern. All specimens, which had been kept deep-frozen after the animals death, were thawed and were then prepared and tested wet, at room temperature (approximately 16 C). Tensile specimens The central section, of uniform cross-sectional shape, was 13 mm long, with a square cross section of 1.8 mm1.8 mm. Youngs modulus of elasticity in bending was determined quasi-statically, in three-point loading, in an Instron 1122 table testing machine, allowance being made for machine

2496 J. D. CURREY
Table 1. The provenance and numbers of the specimens used in the bending tests
Number Species Alligator Axis deer Mother Foetus Bovine 9 year 2 year Crocodile Dolphin Donkey 34 year Dugong Bone Femur Tibia Tibia Metacarpus Tibia Femur Femur Frontal Prefrontal Radius Radius Radius Ulna Scapula Tusk Tibia Radius Bulla Tarso-metatarsus Tibiotarsus Femur Fibula Tibia Humerus BS 26 3 8 9 11 10 20 2 2 2 7 2 2 12 3 9 3 9 5 3 4 5 2 7 [Ca] 26 9 11 10 2 1 2 7 2 2 12 3 9 3 9 5 3 4 5 2 7 p 1 1 3 1 1 2 2 2 2 7 2 9 5 3 2 1 Species Grey seal Horse 7 year Jackass penguin Bone Tibia Femur Humerus Ulna Radius Femur Tibia Humerus Ulna Antler Antler Antler Femur Tarsometatarsus Ossied tendon Tibiotarsus Radius Humerus Femur Humerus Tarsal Radius Tibia Ulna Rib BS 5 9 1 2 2 2 2 9 2 7 25 6 7 13 1 9 6 4 19 2 10 4 24 4 8 Number [Ca] 5 9 1 2 2 2 2 9 2 7 25 6 7 4 1 14 2 7 2 13 2 8 p 2 2 1 2 1 1 2 2 2 4 1 8 2 7 1 7 2 4

King penguin Muntjac deer Red deer Reindeer Roe deer Sarus crane

Elephant Fallow deer Fin whale Flamingo Galapagos tortoise 106 year

Swan Wallaby

White-sided dolphin

Alligator, Alligator mississippiensis; axis deer, Axis axis; bovine, Bos taurus; crocodile, Crocodylus sp.; dolphin, Delphinus sp.; donkey, Equus caballus; dugong, Dugong dugon; elephant, Loxodonta africana; fallow deer, Dama dama; n whale, Balaenoptera physalus; amingo, Phoenicopterus ruber; Galapagos tortoise, Geochelone midas; grey seal, Halichoerus grypus; horse, Equus caballus; jackass penguin, Spheniscus demersus; king penguin, Aptenodytes patagonica; muntjac deer, Muntiacus muntjak; red deer, Cervus elaphus; reindeer, Rangifer tarandus; roe deer, Capreolus capreolus; sarus crane, Grus antigone; swan, Cygnus olor; wallaby, Protemnodon rufogrisea; white-sided dolphin, Lagenorhyncus acutus. BS, number of specimens for which values of bending strength and Youngs modulus of elasticity were determined; [Ca], number of these specimens for which calcium content was also determined; p, number of these specimens for which porosity was also determined; , no values determined.

compliance. An extensometer was then attached to the uniform part of the section of the specimen, which was then loaded in tension at a strain rate of about 0.2 s1. Data were reduced from photographs of the screen of a storage oscilloscope. More recently, the outputs from the load cell and the extensometer were captured and analysed using DASYLAB software. The following mechanical properties were determined for the tensile specimens: Youngs modulus of elasticity (E, actually determined in bending), yield stress (y), yield strain (y) and strain at failure (ult). The yield point (Fig. 1A) was taken to be the point where the curve had deviated by a strain of 0.002 from the straight line describing the initial part of the curve (Currey, 1990). Bending specimens The specimens tested in bending initially all had the same

dimensions (gauge length 30 mm, depth 2 mm, breadth 3.5 mm). Youngs modulus was determined as for the tensile specimens (Fig. 1B). The specimens were broken in threepoint bending, and Youngs modulus and bending strength were calculated using standard beam formulae. The head speed was set so that failure occurred in approximately 2030 s. Other measures The porosity of the cross section approximately 2 mm behind the fracture line was determined for 370 specimens, of which 86 were bending specimens. Porosity was determined by a point-counting method. The mineral content was determined by a colorimetric method and is expressed as milligrams calcium per gram dry defatted bone. Details of the methods are given in Currey (1988).

Bone bending strength 2497 A


30 Yield region Youngs modulus (GPa)




0 Tension Deformation 40 150 200 250 [Calcium] (mg g-1) 300 350

30 Youngs modulus (GPa)



10 Bending Deformation Fig. 1. (A) Idealised load versus deformation curve of a specimen loaded in tension. There is a quite sharply dened yield region. (B) Idealised load versus deformation curve of a specimen loaded in bending. The Youngs modulus of elasticity is calculated from the initial, straight part of the curve using standard beam formulae. The bending strength is calculated from the highest load. 0 150 200 250 300 [Calcium] (mg g-1) 350

Fig. 2. (A) Youngs modulus of elasticity in bending versus calcium content for all specimens. (B) Youngs modulus of elasticity in bending versus calcium content for those specimens whose porosity was measured. Open circles, porosity <8 %; lled circles, porosity >8 %. The value of 8 % was chosen arbitrarily.

Statistical analyses Most of the analyses were least-squares linear regressions on log-transformed data. The data were transformed to reduce the heteroscedasticity and curvilinearity of the plots, and also to make various statistical comparisons more straightforward. However, the raw untransformed data are generally presented here to make the particular form of the relationships (if any) more apparent.

Results Youngs modulus There is clearly a strong relationship calcium content and Youngs modulus (Fig. 2A). The relationship appears as if it might be sigmoidal, but the use of using quadratic and cubic values of calcium content

2498 J. D. CURREY
Table 2. Relationship between Youngs modulus E (GPa), calcium content [Ca] (mg g1) and porosity (%) determined by linear regression
Equation logE=8.58+4.05log[Ca] logE=9.53+4.28log[Ca] logE=1.670.63logp logE=5.06+2.75log[Ca]0.451logp N 855 370 370 370 t 30.35 16.45 18.56 11.48, 13.75 r2 0.52 0.42 0.48 0.62 P <0.001 <0.001 <0.001 <0.001, <0.001

There are two equations for Youngs modulus as a function of calcium content alone. The rst is from the complete data set (see Fig. 2A), the second is from the subset including only those specimens for which values for porosity (p) were measured (see Fig. 2B). In all tables describing the results of statistical analysis, r2 is calculated allowing for the number of degrees of freedom.

as explanatory variables barely improves the statistical t over the use of linear regression. The highest values of calcium content are associated with high moduli, and low values with very low moduli. However, particularly in the region between 230 and 280 mg calcium g1 bone, there is a very large range of values for Youngs modulus. Fig. 2B shows the subset of specimens for which values of porosity were also obtained. The great majority of specimens with low calcium content (<220 mg g1) have a high porosity (>8 %, lled circles), and those that have a high calcium content (>280 mg g1) have a low porosity (open circles). In the middle region, in general, lower values of Youngs modulus are associated with porosities of 8 % or greater. This is borne out by the statistical analysis in which porosity was considered as a continuous variable (Table 2), which shows that 40 % or more of the variance in Youngs modulus can be explained, statistically, by calcium content, and nearly
350 300 250 200 150 A 100 50 0 150 200 250 300 [Calcium] (mg g-1) 350

50 % by porosity. If these two explanatory variables are combined, over 60 % is explained. However, this means that nearly 40 % of the variance is still unexplained by these two variables. It is clear from these equations that porosity and calcium content are themselves related: their correlation coefficients are 0.48 and 0.44 for the raw and logtransformed values, respectively. These coefficients are both highly signicant (P<0.001), though not strong. Bending strength Fig. 3A shows the relationship between calcium content and bending strength (BS). Excluding group A (see below), there is a positive relationship between calcium content and bending strength. Low calcium content is always associated with low bending strength, whereas above approximately 230 mg calcium g1 bone bending strength may be high or low. High-porosity specimens tend to have low bending strengths

300 250 200 150 100 50 0 150 200 250 [Calcium] (mg g-1) 300 350

Bending strength (MPa)

Fig. 3. (A) Bending strength versus calcium content for all specimens loaded to failure in bending. The sample size is smaller than in Fig. 2A because many specimens were not tested to failure in bending. The arrows indicate the subgroup A, consisting of highly mineralised bone from whales, which was excluded from the statistical analysis. (B) Bending strength versus calcium content for the specimens whose porosity was measured. Open circles, porosity <8 %; lled circles, porosity >8 %. Subgroup A is not shown, the porosity of these specimens was <8 %.

Bending strength (MPa)

Bone bending strength 2499

Table 3. Regression relationships between bending strength BS (MPa), calcium content [Ca] (mg g1) and porosity (%)
Equation logBS=4.25+2.70log[Ca] logBS=5.76+3.32log[Ca] logBS=0.39+1.25log[Ca]0.491logp N 262 86 86 t 10.30 4.92 2.05, 7.06 r2 0.29 0.22 0.50 P <0.001 <0.001 <0.05, <0.001

There are two equations for BS as a function of [Ca]; the rst is from the complete data set (Fig. 3A), the second is from the subset including only those specimens for which porosity (p) was measured (Fig. 3B).

Table 4. Regression relationships between bending strength BS (MPa), porosity (%) and Youngs modulus E (GPa)
Equation logBS=1.25+0.851logE logBS=1.18+0.890logE logBS=1.28+0.847E0.053logp N 343 86 86 t 55.46 31.63 21.91, 1.60 r2 0.90 0.92 0.90 P <0.001 <0.001 <0.001, NS

There are two equations for BS as a function of E: the rst is from the complete data set (Fig. 4A); the second is from the subset including only those specimens for which porosity (p) was measured (Fig. 4B). NS, not signicant.

(Fig. 3B). The relationships, though highly signicant (Table 3) are not close ones (r2=0.22, 0.29). Fig. 4A shows the relationship between Youngs modulus and bending strength. Excluding group A, there is an extremely tight, almost linear relationship. Fig. 4B shows that, although specimens with high porosity predominate at the lower end of the distribution, the few high-porosity specimens elsewhere are part of the main distribution. The statistical analysis (Table 4) shows that adding porosity as an explanatory variable had

virtually no effect on the strength of the relationship between logBS and logE. The fact that bending strength (excluding group A) has such a close and nearly proportional relationship with Youngs modulus, with the effect of porosity being very small, immediately suggests that bending strength is determined by some characteristic strain. The initial part of the loaddeformation curve of a bone specimen loaded in bending is essentially straight (Fig. 1B). Since, in these circumstances,

300 250 200 150 100 50 0 0 10 20 Youngs modulus (GPa) 30 40 A 300 250 200 150 100 50 0 0

Bending strength (MPa)

Bending strength (MPa)


20 Youngs modulus (GPa)



Fig. 4. (A) Bending strength versus Youngs modulus of elasticity for all specimens. (B) Bending strength versus Youngs modulus of elasticity for those specimens whose porosity was measured. Open circles, porosity <8 %; lled circles, porosity >8 %. Subgroup A is not shown, the porosity of these specimens was <8 %.

2500 J. D. CURREY
Youngs modulus is equal to stress divided by strain (E=/), if all bones were to fail in bending at the same strain in the outermost bre of the specimen (failure strain ult=k), then bending strength would be exactly proportional to Youngs modulus (=kE). In fact, the equations in Table 4 show that bending strength is not directly proportional to Youngs modulus: for the complete data set, the equation in Table 4 can be rewritten as: BS=15.1E0.85. The fact that the exponent is less than unity suggests that, if bending specimens do fail at some characteristic strain, then that strain is not quite constant, but decreases as a function of Youngs modulus. Unfortunately, it is not possible to determine the yield strain of bending specimens, the onset of yield being too gentle (Fig. 1B). However, the yield strain of tensile specimens can be measured. Such measurements were taken from broadly the same bones as those used to produce the bending data set. The only two tensile specimens from the whales bulla, which had very low strains at yield (0.0017 and 0.0018), were excluded from the statistical analyses. There is a weak, although highly signicant, negative correlation between yield strain y and Youngs modulus E (Fig. 5). The equation is: y = 0.0086 0.000089E , (1) where E is measured in GPa (P<0.001, r2=0.20). If we assume that this equation applies to the bending specimens as well as to the tensile specimens, we can, given the Youngs modulus of the bending specimens, use it to predict their yield strain. It was argued above that, if strain at yield (and therefore failure) is invariant, bending strength should simply be proportional to Youngs modulus. However, Fig. 5 shows that strain at yield is to some extent a negative
0.015 300 250 200 150 100 50 0 0 0 10 20 30 Youngs modulus (GPa) Fig. 5. Strain at yield versus Youngs modulus of elasticity for all tensile specimens. Fig. 6. Bending strength versus yield stress for all specimens. Subgroup A is not shown in this diagram. The broken line is the line of equality. 0 50 100 Yield stress (MPa) 150 200

function of E. Assuming, for the moment, that bone is elasticperfectly brittle and fails at yield, then Ey is the yield stress y and should be equivalent to the bending strength. This yield stress y, expressed in units of MPa, is used below rather than the constant strain implied by the use of Youngs modulus. Fig. 6 shows the relationship between bending strength and yield stress; results from the statistical analysis are given in Table 5. Using yield stress produces only a slightly better t, in terms of r2, than using Youngs modulus (r2=0.90 versus r2=0.88, respectively). The former equation (BS=1.19+1.66y) will give nearly zero bending strength when Youngs modulus is zero, which is satisfactory, whereas the latter equation (BS=32.7+9.55E) gives a large, signicant positive value for bending strength, which must be wrong (Fig. 4; Table 4). Examination of the residuals for the two equations shows the differences. The standardised residuals (expressed in terms of the standard deviation) of yield stress, despite possibly being somewhat heteroscedastic, are much better behaved than the residuals for Youngs modulus, their general distribution being horizontal (Fig. 7B), whereas the residuals for Youngs modulus appear to increase and then decrease as a function of the tted values (Fig. 7A). This was conrmed by regression analysis. Bending strength was tted by linear equations and also by quadratic equations (Table 5). The quadratic equation involving Youngs modulus improved the t slightly, reducing the unexplained variance by 13 %, and also made the curve pass very close to the origin, which is a further improvement. The quadratic equation involving yield stress did not improve the t at all, indicating that the yield stress model is a better one than the Youngs modulus model.


0.010 Yield strain


Bending strength (MPa)

Bone bending strength 2501

Table 5. Regression relationships between bending strength BS (MPa), Youngs modulus E (GPa) and yield stress y (MPa), using quadratic terms
Equation BS=32.7+9.55E BS=2.17+14.8 E0.165E2 BS=1.19+1.66y BS=4.41+1.79y0.0006y2 N 343 343 343 343 t 50.84 19.50, 7.12 54.92 11.31, 0.82 r2 0.88 0.90 0.90 0.90 P <0.001 <0.001, <0.001 <0.001 <0.001, NS

Three of the values of R2 in this table are the same. This is correct, and not caused by typos. NS, not signicant.

Discussion The bending strength of compact bone can best be explained as a function of yield strain; yield strain is similar for all bone, although it declines slightly as Youngs modulus increases. Nevertheless, the bending strength of bone cannot be explained simply by assuming that the bone yields at some strain that is a negative function of Youngs modulus and that the bending moment at failure is predicted by that strain. Fig. 6 shows that although yield stress is directly linearly proportional to bending strength, if bending strength is assumed to equal yield stress, the predicted values of bending strength are too low by a factor of approximately 40 % (Fig. 6, broken line). Bending strength is simple to measure, but this simplicity is misleading. When a specimen is bent, one side is loaded in compression, the other in tension, and strain varies continuously, and in theory linearly, with distance from the neutral axis. If a material is homogeneous and behaves linearly elastically, the stress also varies linearly. Bending strength formulae make this assumption. In an extremely brittle

specimen, the beam formula may be adequate. However, if the outermost bres of a specimen yield, and show some post-yield deformation, the proportionality of stress and strain ceases. The specimen may undergo a greater and greater bending moment, with a resulting larger and larger calculated maximum bending stress, which may be spurious. This matter was analysed theoretically and experimentally for bone by Burstein et al. (1972). Because the stress at a particular strain is nearly proportional to Youngs modulus, for a similar yield strain, a high Youngs modulus will be associated with a high yield stress and therefore a high bending moment when the specimen yields. If, furthermore, the material shows a reasonable amount of post-yield strain, then the bending moment will continue to increase for a while, and the apparent bending strength will be higher. If the bone has a low modulus, it will yield at a rather low stress, and even the large amount of post-yield deformation characteristically shown by low-modulus specimens will not increase the apparent bending strength

3 Residual of Young's modulus 2 1 0 -1 -2 -3 -4 0 100 200 Fitted value 300 400 Residual of yield stress 3 2 1 0 -1 -2 -3 -4 0


200 Fitted value



Fig. 7. (A) Standardised residuals (expressed in terms of their standard deviations) versus tted values in the equation BS=32.7+9.55E, where BS is bending strength and E is Youngs modulus of elasticity. (B) Standardised residuals versus tted values in the equation BS=1.19+1.66y, where y is yield stress.

2502 J. D. CURREY
350 300 10 Bending strength (MPa) 250 200 150 100 A 50 1 0 0 5 10 15 20 25 30 35 0 50 100 150 200 250 300 350 Youngs modulus (GPa) Fig. 8. Strain ratio (ultimate strain/yield strain) versus Youngs modulus of elasticity for tensile specimens. Note that the ordinate is on a logarithmic scale. Predicted bending strength (MPa) Fig. 9. Relationship between observed bending strength and that predicted from the theory of Burstein et al. (1972) and regressions from Figs 5 and 8. The isolated whale bulla specimens labelled A do not form part of the general pattern. The broken line is the line of equality.

Strain ratio

sufficiently to make up for the low stress at yield. Burstein et al. (1972) showed that, for a rectangular cross section, as was used in the present tests, if the post-yield strain is equal to the elastic strain, then the bending moment will be raised by a factor of approximately 1.5 compared with a completely brittle material. If the post-yield strain is ve times the elastic strain, then the factor is 1.7. It requires, therefore, little post-yield strain to reap most of the benets of not being completely brittle. This post-yield behaviour of bone may partly explain why the predictions of yield stress in the present study, which assume that bone is linearly elastictotally brittle, are too low. The ratio of ultimate strain ult to yield strain y, or strain ratio ult/y, is shown in Fig. 8 plotted against Youngs modulus for all the tension specimens. The equation for the relationship is: log(ult/y) = 0.996 0.0289logE (2)

(P<0.001, r2=0.37). The ordinate is plotted on a logarithmic scale; strain ratios near unity indicate brittle or near-brittle behaviour. Many of the more compliant specimens have a strain ratio of the order of 1015. According to the calculations of Burstein et al. (1972), such specimens would be expected to have a bending strength approximately twice their yield strength. In the present specimens, therefore, there is a variation of approximately a factor of two in the difference between the yield strength and the bending strength; the bones with a lower Youngs modulus tend to have higher factors (Fig. 8). This phenomenon of post-yield resistance to bending would, therefore, seem to be of the right magnitude to explain why the bending strengths of the specimens are greater than

those predicted by yield stress. It is surprising, however, that some of the more brittle bones do not show a lower bending strength, closer to their presumed yield stress. It may be taking theory too far to apply the equations of Burstein et al. (1972) to the present data because to do so requires the building of prediction on prediction. It requires the combination of two regressions: yield strain as a function of Youngs modulus and strain ratio as a function of Youngs modulus. Both these relationships are rather loose. The calculations of Burstein et al. (1972) use , dened as the ratio of pre-yield strain to ultimate strain (the inverse of the strain ratio), where C=2{1[1(1)2]/(1)2}. The ratio of calculated bending stress to stress at yield in tension is then given by 0.25{C(32)+2(2C)3/C}. These calculations assume that the specimens have a rectangular cross section (which is the case in this investigation) and that the specimens do not yield in compression before failure. If the specimens do yield in compression, the effect will be negligible down to a value of of approximately 0.5. For lower values of , the calculated value of bending strength starts to deviate less from the actual yield stress than predicted by the above equations. The value of as a function of Youngs modulus was calculated from the regression equation for the data shown in Fig. 8, and the strain at yield as a function of Youngs modulus was calculated from the regression equation for the data shown in Fig. 5. The results of these calculations are shown in Fig. 9. Except for the group of eight outliers that form subgroup A (see

Bone bending strength 2503

below), the predicted values and the actual values are closely similar until a predicted value of bending strength of approximately 180 MPa. Thereafter, the actual values become greater than the predicted values. Another factor that may be important for the present results was rst analysed by Weibull (1951). A tensile specimen undergoes roughly the same stress all through its volume, while a bending specimen undergoes high tensile stresses within a rather small volume close to one surface opposite the central loading point. Any real material is not quite uniform and has a distribution of strengths throughout its volume. It is likely, therefore, that a tensile specimen will yield at a lower calculated stress than a bending specimen simply because the stress in a tensile specimen has a larger volume over which to seek out a weak part of the specimen. Therefore, the stresses in the outermost bres of the bending specimens before they yield will on average be higher than the overall yield stress reached if the same specimens had been loaded in tension. The values used to calculate yield stress here were from tensile specimens, which may explain why the bending moments of the bending specimens tended to be higher than predicted by the yield stress model. The specimens that deviated most were the stronger specimens, which had the highest Youngs modulus and were, in general, more brittle. The Weibull effect is more pronounced in brittle materials; however, its magnitude cannot be quantied from the present data. The specimens that comprised subgroup A (Figs 3, 4, 9) were from very highly mineralised tympanic bullae from the n whale Balaenoptera physalus (Currey, 1979) with a high Youngs modulus. They had a low bending strength (Figs 3, 4), a low strength in tension (approximately 25 MPa) and a very low strain at yield and fracture. These specimens were completely brittle, showing no post-yield strain in tension. Also, in tension, they broke at a lower strain than all the other bones. Therefore, in bending, they cannot make use of their high Youngs modulus, both because they break at a lower strain than other bones and because they undergo no post-yield deformation, so the bending moment cannot increase after yield. In this very highly mineralised bone, the few cracks that appear in the pre-yield region presumably nd it so easy to travel that they become fatal, giving a strain at yield (approximately 0.002) considerably less than that reached by other bones (>0.004), and the material fails in a brittle manner at a low bending moment. Professor Mike Ashby, Miss Debra Balderson and Drs Terry Crawford and Justin Molloy gave me helpful advice. I thank an anonymous referee who encouraged me actually to apply the predictions of Burstein et al. (1972) to the data. The results were surprising and interesting.

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