Interspecific Scaling Research Articles

Scaling Relationship between Body Weight and Fermentation Gut Capacity in Axis Deer

The scaling relationship between body weight and fermentation gut capacity presumably impacts dietary patterns of ruminants. Interspecific scaling relationships between body weight and fermentation gut capacity have a slope of 1.0 (isometric scaler) when body weights span � 3 orders of magnitude. Isometric scaling relationships between body weight and fermentation gut capacity also have been suggested to explain dietary patterns within species, but whether scaling relationships are isometric within a single species is unresolved. We determined if the intraspecific scaling relationship between body weight and fermentation gut capacity was isometric for axis deer (Axis axis) and assessed the influence of reproductive status of females on the scaling relationship. We collected 8 adult male and 21 adult female axis deer from January 2004 to April 2005 at a private ranch in Bastrop County, Texas. Capacity of the rumen‐reticulum, cecum, and colon, organs where fermentation occurs, was estimated by calculating wet weight and volume capacity. Forage quality, month that animal was collected, and sex did not influence scaling relationships. Regressions apparently had isometric scalers for fermentation gut capacity but the relationships were not strong, in part because females that were pregnant had smaller fermentative gut capacities than nonreproductive or lactating animals.

Convergence of metabolic rate of cultured cells from animals of different sizes

cultured fibroblasts of cell lines from animals of different size have similar activities for a range of different enzymes and metabolic rates that converge on the similar values despite the metabolic rates of the animals of different sizes having quite distinctly different basal metabolic rates (

Body size and scaling of the hands and feet of prosimian primates

The hands and feet of primates fulfill a variety of biological roles linked with food acquisition and positional behavior. Current explanations of shape differences in cheiridial morphology among prosimians are closely tied to body size differences. Although numerous studies have examined the relationships between body mass and limb morphology in prosimians, no scaling analysis has specifically considered hand and foot dimensions and intrinsic proportions. In this study, we present such an analysis for a sample of 270 skeletal specimens distributed over eight prosimian families. The degree of association between size and shape was assessed using nonparametric correlational techniques, while the relationship between each ray element length and body mass (from published data and a body mass surrogate) was tested for allometric scaling. Since tarsiers and strepsirrhines encompass many taxa of varying degrees of phylogenetic relatedness, effective degrees of freedom were calculated, and comparisons between families were performed to partially address the problem of statistical nonindependence and "phylogenetic inertia." Correlational analyses indicate negative allometry between relative phalangeal length (as reflected by phalangeal indices) and body mass, except for the pollex and hallux. Thus, as size increases, there is a significant decrease in the relative length of the digits when considering all prosimian taxa sampled. Regression analyses show that while the digital portion of the rays scales isometrically with body mass, the palmar/plantar portion of the rays often scales with positive allometry. Some but not all of these broadly interspecific allometric patterns remain statistically significant when effective degrees of freedom are taken into account. As is often the case in interspecific scaling, comparisons within families show different scaling trends in the cheiridia than those seen across families (i.e., lorisids, indriids, and lemurids exhibit rather different allometries). The interspecific pattern of positive allometry that appears to best characterize the metapodials of prosimians, especially those of the foot, parallels differences found in the morphology of the volar skin. Indeed, relatively longer metapodials appear to covary with flatter and more coalesced volar pads, which in turn slightly improve frictional force for animals that are at a comparative disadvantage while climbing because of their larger mass. Despite the essentially isometric relationship found between digit length and body mass across prosimians, examination of the residual variation reveals that tarsiers and Daubentonia possess, relative to their body sizes, remarkably long fingers. Such marked departures between body size and finger length observed in these particular primates are closely linked with specialized modes of prey acquisition and manipulation involving the hands.

Interspecific scaling of the morphology and posture of the limbs during the locomotion of cats (Felidae)

For phylogenetically diverse mammals, ranging from small rodents to large ungulates, the generalization that limb erectness increases with increased size is supported by some size-dependent scaling relationships of appendicular skeletal anatomy as well as a limited number of direct observations of limb posture during locomotion. If size alone is the causal basis for different limb posture, then the erectness of limbs should increase significantly with increased size within a phylogenetically narrow lineage, but such data are sparse. Thus, to better establish the correlation between size and posture of mammalian limbs, we quantified the scaling relationships between mass and limb dimensions and kinematics during walking of nine species within the felid (cat) clade, which has qualitatively similar limb design. We studied the domestic cat, serval, ocelot, lynx, leopard, cheetah, cougar, lion and tiger, which had masses ranging from <4 kg to nearly 200 kg. Apart from variation associated with overall size, the lengths of the appendicular skeletal structures of most of the felid species were morphologically very similar in multivariate space. The kinematics of the limbs were also relatively uniform, and size had little predictive value for limb posture among felid species. Only three out of a total of 24 angular variables at footfall and midstance changed significantly (0.02<P<0.05) with increased mass. Thus, in contrast to previous broadly comparative studies of mammals, larger species of felids did not have more upright limbs than smaller species.

Morphological and biochemical jumping adaptations in smaller grasshoppers

Many interspecific vertebrate studies have shown that similarly shaped animals have isometric proportions of muscle mass and leg length, and equivalent jumping performance. In contrast, ontogenetic grasshopper studies have shown that older grasshoppers have disproportionally longer legs, produce more lactate during jumping, and jump more body lengths. In this study, we compared the morphology, jump performance and lactate production of twenty species of female grasshoppers ranging in size from 0.2 to 7.0 g to determine whether interspecific scaling of jump performance followed vertebrate or ontogenetic grasshopper models. In contrast to both vertebrate and ontogenetic locust studies, we found that smaller grasshoppers had a 6-fold greater proportion of muscle mass and relatively longer hind legs. Although these morphological changes should result in improved performance, we found that body size did not affect jump distance or jump energy (either maximally or throughout the ten minute trial). Unlike the ontogeny study, smaller grasshoppers produced more lactate during jumping. Model calculations suggest that increased leg length, lactate production and muscle size are necessary for smaller grasshoppers to overcome a greater cost of drag and yet achieve similar jump distances. This work was partially supported by NSF IBN-9985857 and IOB-0419704 to JFH and IBN-0104959 to SDK and JFH.

Lumbar ontogenetic allometry and dimorphism in humans. A case for comparison between interspecific and intraspecific scaling

The ontogenetic allometry of the lumbar region of 1913 humans (1228 females and 685 males), ranging from newborn to 21-year-old individuals, was studied by means of length, width, projected surface area and bone mineral density of the segment L2 - L4, obtained by dual X-ray absorptiometry (DXA). All these parameters were regressed to body mass and height of the individuals, considered alternatively as the independent variable. Firstly, we addressed the comparison between the results obtained on both sexes in order to elucidate whether ontogenetic differences existed. Length of the segments increased significantly faster in females than in males, independently whether the regression was made against body mass or height, while in both types of regression width scaled in males faster than in females. Regarding bone mineral density, although males increased bone mineral density faster than females, slope differences were not significant. However, y-interception was significantly higher in females than in males when bone mineral density was regressed to body mass. Results on length and width are compared with others from previous research on allometry. Finally, global results are discussed as regards the slope predictions for interspecific scaling.

Invariant scaling of phytoplankton abundance and cell size in contrasting marine environments

Scaling relationships such as the variation of population abundance with body size provide links between individual organisms and ecosystem functioning. Previous work, in marine pelagic ecosystems, has focused on the relationship between total phytoplankton abundance and the assemblage mean cell size. However, the relationship between specific population abundance and cell size in marine phytoplankton has received little attention. Here, we show that cell size accounts for a significant amount of variability in the population abundance of phytoplankton species across a cell volume range spanning seven orders of magnitude. The interspecific scaling of population abundance and cell size takes a power exponent near -3/4. Unexpectedly, despite the constraints imposed on large phytoplankton by limited resource acquisition, the size scaling exponent does not differ between contrasting marine environments such as coastal and subtropical regions. These findings highlight the adaptive abilities of individual species to cope with different environmental conditions and suggest that a general rule such as the 'energetic equivalence' constrains the abundance of phytoplankton populations in marine pelagic ecosystems.

Some mathematical and technical issues in the measurement and interpretation of open-circuit indirect calorimetry in small animals

Indirect calorimetry is increasingly used to investigate why compounds or genetic manipulations affect body weight or composition in small animals. This review introduces the principles of indirect (primarily open-circuit) calorimetry and explains some common misunderstandings. It is not widely understood that in open-circuit systems in which carbon dioxide (CO2) is not removed from the air leaving the respiratory chamber, measurement of airflow out of the chamber and its oxygen (O2) content paradoxically allows a more reliable estimate of energy expenditure (EE) than of O2 consumption. If the CO2 content of the exiting air is also measured, both O2 consumption and CO2 production, and hence respiratory quotient (RQ), can be calculated. Respiratory quotient coupled with nitrogen excretion allows the calculation of the relative combustion of the macronutrients only if measurements are over a period where interconversions of macronutrients that alter their pool sizes can be ignored. Changes in rates of O2 consumption and CO2 production are not instantly reflected in changes in the concentrations of O2 and CO2 in the air leaving the respiratory chamber. Consequently, unless air-flow is high and chamber size is small, or rates of change of O2 and CO2 concentrations are included in the calculations, maxima and minima are underestimated and will appear later than their real times. It is widely appreciated that bigger animals with more body tissue will expend more energy than smaller animals. A major issue is how to compare animals correcting for such differences in body size. Comparison of the EE or O2 consumption per gram body weight of lean and obese animals is misleading because tissues vary in their energy requirements or in how they influence EE in other ways. Moreover, the contribution of fat to EE is lower than that of lean tissue. Use of metabolic mass for normalisation, based on interspecific scaling exponents (0.75 or 0.66), is similarly flawed. It is best to use analysis of covariance to determine the relationship of EE to body mass or fat-free mass within each group, and then test whether this relationship differs between groups.

Beyond the ‘3/4‐power law’: variation in the intra‐and interspecific scaling of metabolic rate in animals

In this review I show that the '3/4-power scaling law' of metabolic rate is not universal, either within or among animal species. Significant variation in the scaling of metabolic rate with body mass is described mainly for animals, but also for unicells and plants. Much of this variation, which can be related to taxonomic, physiological, and/or environmental differences, is not adequately explained by existing theoretical models, which are also reviewed. As a result, synthetic explanatory schemes based on multiple boundary constraints and on the scaling of multiple energy-using processes are advocated. It is also stressed that a complete understanding of metabolic scaling will require the identification of both proximate (functional) and ultimate (evolutionary) causes. Four major types of intraspecific metabolic scaling with body mass are recognized [based on the power function R=aMb, where R is respiration (metabolic) rate, a is a constant, M is body mass, and b is the scaling exponent]: Type I: linear, negatively allometric (b<1); Type II: linear, isometric (b=1); Type III: nonlinear, ontogenetic shift from isometric (b=1), or nearly isometric, to negatively allometric (b<1); and Type IV: nonlinear, ontogenetic shift from positively allometric (b>1) to one or two later phases of negative allometry (b<1). Ontogenetic changes in the metabolic intensity of four component processes (i.e. growth, reproduction, locomotion, and heat production) appear to be important in these different patterns of metabolic scaling. These changes may, in turn, be shaped by age (size)-specific patterns of mortality. In addition, major differences in interspecific metabolic scaling are described, especially with respect to mode of temperature regulation, body-size range, and activity level. A 'metabolic-level boundaries hypothesis' focusing on two major constraints (surface-area limits on resource/waste exchange processes and mass/volume limits on power production) can explain much, but not all of this variation. My analysis indicates that further empirical and theoretical work is needed to understand fully the physiological and ecological bases for the considerable variation in metabolic scaling that is observed both within and among species. Recommended approaches for doing this are discussed. I conclude that the scaling of metabolism is not the simple result of a physical law, but rather appears to be the more complex result of diverse adaptations evolved in the context of both physico-chemical and ecological constraints.

Lumbar ontogenetic allometry and dimorphism in humans. A case for comparison between interspecific and intraspecific scaling

Lumbar ontogenetic allometry and dimorphism in humans. A case for comparison between interspecific and intraspecific scaling

Biomechanical consequences of scaling

To function over a lifetime of use, materials and structures must be designed to have sufficient factors of safety to avoid failure. Vertebrates are generally built from materials having similar properties. Safety factors are most commonly calculated based on the ratio of a structure's failure stress to its peak operating stress. However, yield stress is a more likely limit, and work of fracture relative to energy absorption is likely the most relevant measure of a structure's safety factor, particularly under impact loading conditions characteristic of locomotion. Yet, it is also the most difficult to obtain. For repeated loading, fatigue damage and eventual failure may be critical to the design of biological structures and will result in lower safety factors. Although area:volume scaling predicts that stresses will increase with size, interspecific comparisons of mammals and birds show that skeletal allometry is modest, with most groups scaling (l proportional, variant d0.89) closer to geometric similarity (isometry: l proportional, variant d1.0) than to elastic similarity (l proportional, variant d0.67) or stress similarity (l proportional, variant d0.5). To maintain similar peak bone and muscle stresses, terrestrial mammals change posture when running, with larger mammals becoming more erect. More erect limbs increases their limb muscle mechanical advantage (EMA) or ratio of ground impulse to muscle impulse (r/R= integral G/integral Fm). The increase in limb EMA with body weight (proportional, variant W0.25) allows larger mammals to match changes in bone and muscle area (proportional, variant W0.72-0.80) to changes in muscle force generating requirements (proportional, variantW0.75), keeping bone and muscle stresses fairly constant across a size range 0.04-300 kg. Above this size, extremely large mammals exhibit more pronounced skeletal allometry and reduced locomotor ability. Patterns of ontogenetic scaling during skeletal growth need not follow broader interspecific scaling patterns. Instead, negative allometric growth (becoming more slender) is often observed and may relate to maturation of the skeleton's properties or the need for younger animals to move at faster speeds compared with adults. In contrast to bone and muscle stress patterns, selection for uniform safety factors in tendons does not appear to occur. In addition to providing elastic energy savings, tendons transmit force for control of motion of more distal limb segments. Their role in elastic savings requires that some tendons operate at high stresses (and strains), which compromises their safety factor. Other 'low stress' tendons have larger safety factors, indicating that their primary design is for stiffness to reduce the amount of stretch that their muscles must overcome when contracting to control movement.

Testing models of biological scaling with mammalian population densities

Two hypotheses have been suggested to explain the form of interspecific scaling of organismal characteristics to body size, such as the well-known increase in total metabolism with body mass. A hypothesis based on simple Euclidean geometry suggests that the scaling of many biological variables to body size should have a scaling exponent of 2/3, or [Formula: see text]0.667. On the other hand, according to a hypothesis based on fractal dimensions, the relationship between biological variables and body mass should have a scaling exponent of 0.750. We conducted a power analysis of the predicted exponents of scaling under the Euclidean and fractal hypotheses, using average adult body masses and population densities collected from the published literature on mammalian species. The collected data reflect 987 mammal populations from a broad variety of terrestrial habitats. Using statistical methods we determined the sample sizes required to decide between the values of the scaling exponent of the density-to-mass relationship based on the Euclidean (–0.667) and fractal (–0.750) hypotheses. Non-linearities in the dataset and insufficient power plagued our tests of the predictions. We found that mammalian species weighing less than 100 kg had a linear scaling pattern, sufficient power to reveal a difference between the scaling coefficients –0.667 and –0.750, and an actual scaling coefficient of –0.719 (barely significantly different from –0.667 but not from –0.750). Thus, our results support the fractal hypothesis, though the support was not particularly strong, which suggests that the relationship between body mass and population density should have a scaling exponent of –0.750.

Global allocation rules for patterns of biomass partitioning in seed plants.

A general allometric model has been derived to predict intraspecific and interspecific scaling relationships among seed plant leaf, stem, and root biomass. Analysis of a large compendium of standing organ biomass sampled across a broad sampling of taxa inhabiting diverse ecological habitats supports the relations predicted by the model and defines the boundary conditions for above- and below-ground biomass partitioning. These canonical biomass relations are insensitive to phyletic affiliation (conifers versus angiosperms) and variation in averaged local environmental conditions. The model thus identifies and defines the limits that have guided the diversification of seed plant biomass allocation strategies.

Comparing interspecific and intraspecific allometry in the Anatidae

Interspecific scaling relationships (e.g. of limb size with body mass) in vertebrates are usually assumed to be functional (e.g. biomechanical) attributes. In this paper on the Anatidae, we study the scaling of wing length and tarsus length with body mass, relationships that can be expected to have biomechanical significance. At an interspecific level, both wing length and tarsus are positively allometric, a finding consistent with results from previous comparative avian studies. These trends remained significant in regressions controlling for the effects of phylogeny, but interspecific slopes were less steep within tribes than in the whole family (a taxon-level effect). We are not aware of any biomechanical resasons that explain these patterns satisfactorily. Intraspecific (static) allometries in Green-winged Teal (Anas crecca) and Marbled Teal (Marmaronetta angustirostris) are different: wing length is negatively allometric and tarsus is isometric. These anomalies suggest that interspecific and intraspecific scaling relationships do not share common causes. Our results bring into question the significance of interspecific allometries in vertebrate morphology, which may to some extent be non-functional by-products of morphological optimisation processes within species and ecological differences between them.

Comparing interspecific and intraspecific allometry in the Anatidae

Interspecific scaling relationships (e.g. of limb size with body mass) in vertebrates are usually assumed to be functional (e.g. biomechanical) attributes. In this paper on the Anatidae, we study the scaling of wing length and tarsus length with body mass, relationships that can be expected to have biomechanical significance. At an interspecific level, both wing length and tarsus are positively allometric, a finding consistent with results from previous comparative avian studies. These trends remained significant in regressions controlling for the effects of phylogeny, but interspecific slopes were less steep within tribes than in the whole family (a taxon-level effect). We are not aware of any biomechanical resasons that explain these patterns satisfactorily. Intraspecific (static) allometries in Green-winged Teal (Anas crecca) and Marbled Teal (Marmaronetta angustirostris) are different: wing length is negatively allometric and tarsus is isometric. These anomalies suggest that interspecific and intraspecific scaling relationships do not share common causes. Our results bring into question the significance of interspecific allometries in vertebrate morphology, which may to some extent be non-functional by-products of morphological optimisation processes within species and ecological differences between them.

Interspecific scaling of the hindlimb skeleton in lizards, crocodilians, felids and canids: does limb bone shape correlate with limb posture?

AbstractDuring locomotion, lizards and crocodilians generally use a more sprawling limb posture than most mammals and experience substantial axial rotation of the femur. Consequently, the limb bones of most mammals are loaded predominantly in bending, but the limb bones of lizards and crocodilians are loaded primarily in torsion. As body size increases, torsional shear stress in limb bones is expected to increase more than bending stress; therefore, limb bone diameters of lizards and crocodilians might be expected to scale with relatively greater positive allometry than limb bone diameters of mammals that use upright posture. To test this hypothesis, scaling patterns of the femur and tibia in lizards (iguanians and varanids) and crocodilians were compared with patterns in felid and canid mammals, using both non‐phylogenetic statistical methods and phylogenetically independent contrasts. Comparisons with theoretical models indicate that size‐related changes in limb bone geometry do not completely compensate for size‐related increases in limb bone stress in the lizard or crocodilian lineages examined. Unless lizards and crocodilians compensate for size‐related increases in limb bone stress through other mechanisms (e.g. changes in limb kinematics or the mechanical properties of limb bones), limb bone stresses are predicted to be relatively greater among larger species of these lineages. However, limb bone diameters appear to scale with greater positive allometry (relative to body mass) in varanids than in iguanians, suggesting that larger lizard lineages might compensate for increased stress through changes in bone geometry to a greater degree than smaller lineages. Allometric scaling patterns for many limb bone diameters among iguanians are more similar to those of felids and canids than to those of varanids; thus, sprawling locomotor habits do not correlate clearly with a particular pattern of limb bone scaling. This suggests that similarity of interspecific scaling patterns of limb bone lengths and diameters is not sufficient to justify inferences of similar locomotor function.

Interspecific scaling of the hindlimb skeleton in lizards, crocodilians, felids and canids: does limb bone shape correlate with limb posture?

Interspecific scaling of the hindlimb skeleton in lizards, crocodilians, felids and canids: does limb bone shape correlate with limb posture?

The size–grain hypothesis and interspecific scaling in ants

1. The size–grain hypothesis maintains that as terrestrial walking organisms decrease in size, their environment becomes less planar and more rugose. The benefits of long legs (efficient, speedy movement over a planar environment) may thus decrease with smaller body size, while the costs (larger cross‐sectional area limiting access to the interstitial environment) are enhanced. 2. A prediction from this hypothesis – that leg size should increase proportionately with body mass – is examined. Ants are among the smallest walking animals and extend the size gradient five orders of magnitude beyond the traditional ‘mouse to elephant’ curve. The mass of 135 species of worker ants spans 3·7 orders of magnitude (0·008–53 mg). Larger ants tended to be slimmer and longer legged. Ant subfamilies varied in their scaling relationships, but four out of five showed a positive allometry for hind leg length (b &gt; 0·33). Mammals, in contrast, show isometry for leg length over six orders of magnitude. 3. It is suggested that ants make a transition from living in an interstitial environment when small to a planar environment when large, a habit continued by most terrestrial mammals. Head length and pronotum width are robust estimators of mass in ants.

Standard metabolic rate of the largest Australian lizard, Varanus giganteus

The intraspecific relationship between standard metabolic rate (VO2; ml hr−1) and body mass for Varanus giganteus is 0.0896 g0.90 at 25°C and 0.126 g0.96 at 35°C. The relationship between VCO2 (ml hr−1) and body mass is 0.052 g0.92 at 25°C and 0.094 g0.97 at 35°C. The common slope for the intraspecific mass exponent at the two body temperatures of 25 and 35°C was 0.93 for VO2 and 0.95 for VCO2. When these data for V. giganteus are considered in conjunction with those from recent studies of other varanids, the interspecific scaling exponent for Varanus is approximately 0.9; this is similar to the intraspecific scaling exponent for V. giganteus and other varanids.

THE SCALING OF PLANT AND ANIMAL BODY MASS, LENGTH, AND DIAMETER.

The interspecific scaling exponents of body mass M and diameter D with respect to length L were determined to evaluate the predictions of three scaling hypotheses (geometric, stress, and elastic similitude). The relation between M and L was determined for data from a total of 133 aquatic and terrestrial species (66 plant and 67 animal species); the relation between D and L was determined independently for a total of 753 aquatic and terrestrial species (667 plant and 86 animal species). Organisms were crudely classified as to their geometry (spheres, spheroids, cylinders) and shape (defined as the body slenderness factor, L/D) to examine whether geometry and shape evinced size-dependent variations. Regression indicated M = 1.29L2.95 (r2 = 0.91, N = 133; αRMA = 3.09 ± 0.088). The stress and elastic similitude (which respectively predict αRMA = 5 and αRMA = 4) were rejected; geometric similitude was not (αRMA = 3). For animals and plants, αRMA = 2.81 ± 0.061 (r2 = 0.98), and αRMA = 2.95 ± 0.093 (r2 = 0.94), respectively. For aquatics and terrestrial organisms, αRMA = 2.82 ±0.134 (r2 = 0.97, N = 36), and αRMA = 3.08 ±0.111 (r2 = 0.89, N = 97), respectively. These results were interpreted to support the hypothesis of geometric similitude. For the pooled plant and animals data, D = 0.05L1.00 (r2 = 0.95; αRMA = 1.03 ± 0.009), which was compatible with the hypothesis of geometric similitude. For plants, D = 0.05L1.06 (r2 = 0.95; αRMA = 1.09). For animals, D = 0.29L0.98 (r2 = 0.95; αRMA = 1.01 ± 0.025). Also, for aquatics, αRMA = 0.951 ± 0.151, whereas for terrestrial plants and animals, αRMA = 1.03 ± 0.089. Although the scaling exponent for D differed among individual groupings of animals and plants, the results of regression analyses were interpreted to indicate that, on the average, body diameter scaled isometrically with respect to length as predicted by geometric similitude. For the pooled data set, organic shape varied over 3 orders of magnitude; L varied over 9 orders of magnitude reflecting 22 orders of magnitude of M. In terms of body geometry and the absolute numbers of species in the total data set: spherical shaped species (L = D) < unassigned species < prolate spheroidal species < cylindrical (squat < slender) species. The largest organisms in the data set were slender (L/D > 20) cylindrical plants; the smallest organisms were spherical plants and animals. Although not subject to statistical inference, these data were interpreted to indicate that organic shape and geometry evince size-dependent variations. These variations as well as size-dependent changes in bulk density are hypothesized to account for the scaling exponents of M and D determined for individual plant and animal clades and grades.