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Intraspecific scaling of flight power in the bat Glossophaga soricina (Phyllostomidae).

Aerodynamic theory predicts that power output during flight should vary with body mass by an exponent of 1.56 when wing morphology remains constant (within an individual), and by an exponent of 1.19 when wing morphology changes with body mass (within a species or between species). I tested these predictions by estimating the power input during horizontal flight in three pregnant and two subadult Glossophaga soricina using a multivariate regression model. This analysis yielded power input during resting and flight as well as the energetic equivalent of change in body mass. A comparison of the estimated flight power for pregnant G. soricina, with published data on flight power of nonpregnant adults, revealed that energy turnover in flight is highest for pregnant G. soricina. Flight power of a 13-g pregnant G. soricina was even higher than that of a 16-g non-pregnant Glossophaga longirostris. A least-squares regression analysis yielded the following equations for the intraspecific scaling of flight power with body mass: power input during horizontal flight (Pf) = 24099 body mass (bm; kg)2.15 (r2 = 0.97) for the intra-individual allometry (pregnancy) and Pf = 113 bm(kg)0.95 (r2 = 0.99) for the inter-individual allometry (ontogeny). Both mass exponents are not significantly different from the predicted values for the scaling relationship of flight power within an individual (1.56) and within a species (1.19). This is the first measurement of power input during flight for subadult and pregnant bats.

Adaptation, Physiological↗

Mandibular growth rates in human fetal development.

A morphometric analysis of changing proportions in the developing mandible was undertaken in 18 human embryos and fetuses of both sexes (developmental age from 8 to 14 weeks, crown-rump length, CRL, from 34 to 110 mm), previously cleared and stained with a specific method for bone (alizarin red S). Reference points were located on the mandible, i.e. condylar process (Pcl), coronoid process (Pco), gnathion (GN), gonion (GO), superior symphyseal point (SSP), for measuring linear dimensions, i.e. Pcl-GN, Pcl-Pco, Pco-GN, GO-GN, SSP-GN. The gonial (Pcl-GO-GN) and the (Pcl-GN-Pcl) angles were also measured. All linear dimensions were correlated with the CRL by bivariate allometry (1n y = 1n a+b 1n x): they all grew with positive allometry, except GO-GN with isometry. The mandibular ramus grew relatively faster than the body, both in length and height, and the greatest growth rate was found for ramus height. The relation between mandibular shape and the craniofacial structures was investigated using scale drawings obtained from photographs of fetal skulls in lateral view. In the youngest fetuses the mandible was prognathic, then became retrognathic. During the period investigated the zygomatic process and squama of the temporal bone were in a lower and more inclined position in relation to the transverse plane passing through the zygomatic arch than in the newborn and adult. This study identifies parameters fitting changing trends in height, length and shape of the human mandible during the prenatal period (8-14 weeks); moreover, it emphasizes that the mandibular growth patterns differ significantly from those of successive development periods.

Adult↗

Allometric nucleo-cytoplasmic volune relations and rotation ellipsoid neuronal bodies of the cat nodosal and superior sympathetic ganglia.

Using a Henning and Elias' plot, evidence was obtained for the rotation ellipsoid nature of the three-dimensioinal shape of neuronal bodies of the nodosal and cervical sympathetic ganglia of the cat. Nucleo-cytoplasmic volume relationships for both types of ganglia neurons obeyed the allometry formula. The allometry coefficient (ratio of logarithms of cytoplasmic and nuclear volumes) was found to be similar in the neurons of both ganglia, its value corresponding to 1.38--1.43.

Animals↗

Extrapolation of animal toxicity data to man.

Laboratory animals are used as models for humans in toxicity studies. This use is based on the assumption that extrapolation of biological data from animals to humans is valid. Three methods of extrapolation are considered: the use of body mass equivalence, caloric scaling across species, and the use of the surface area equivalence. Allometry, defined as the study of size and its consequences, is considered. There is still controversy whether there is an allometric relationship for energy metabolism. Allometry offers, among others, the concept that not all of the mass of the animal is equally involved in metabolism. In recent years the principles of pharmacokinetics have been applied to interspecies scaling; pharmacokinetic short-term studies can be used to determine whether allometric scaling is justified. Considerations, however, should be given to (pharmacokinetic) differences in the same species and to species variability. It would be useful to develop a set of criteria for deciding when the pharmacokinetic model is needed and when simpler models will suffice.

Administration, Inhalation↗

Louis Bolk revisited. II--Retardation, hypermorphosis and body proportions of humans.

A number of morphological particularities of humans, generally thought to be specializations produced by natural selection, arguably are instances of hypermorphosis: alterations in proportion brought about by simple prolongation of general embryonic allometries. The descended larynx, the flattened human chest, and several limb proportions are given as examples. The case of the human foot, often quoted as an example of advanced specialization in humans, is examined in some more detail. Besides, it is argued that the human growth pattern displays a unique 7-year periodicity unlikely to be the result of the erratics of Darwinian evolution. Developmental acceleration is the unavoidable correlate of animal specialization; it brings about the breakdown of the periodicity present in the generalized humanlike development and causes the embryonic allometries to be cut off prematurely in the non-human mammal.

Adult↗

Morphometry of the human heart in the second and third trimesters of gestation.

The growth of the heart was studied in 56 human fetuses ranging from 14 to 38 weeks (23 female and 33 male) using the following measurements: cardiac length and width, aortic and pulmonary artery internal diameters, right and left anterior ventricular wall thickness. Relationships between the fetal cardiac data and crown-rump length were explored by allometry and linear regression analysis. The present study confirmed that the heart increases with high growth rates and positive allometry during prenatal life. Statistically different growth rates were found comparing female with male fetuses (sexual dimorphism): (a) in the third trimester (P < 0.001, except for the cardiac length and the left ventricular wall thickness), (b) considering fetuses of the second and third trimesters together (only for cardiac width, P = 0.0015). In these circumstances the female fetuses had the greatest growth rates. The length and the width of the heart are diameters easily obtained in the ultrasonographic four-chamber view of the heart in the last two trimesters of gestation. Therefore, these data can be analyzed by non-invasive methods and are useful for prenatal detection of congenital heart disease.

Biometry↗

On the assessment of the growth patterns in human fetal limbs: longitudinal measurements and allometric analysis.

The total length (TL) and length of the ossified part (OL) of some long bones of the upper (humerus, ulna, radius) and lower limb (femur, tibia, fibula) were evaluated in 58 aborted human fetuses (crown-rump length, CRL, between 38 and 116 mm, developmental age from 8 to 14 weeks). The specimens, without any detectable malformation, were cleared and double-stained with alcian blue and alizarin red S to obtain a differential detection of the ossified part within the comprehensive outline between the cartilaginous epiphyses. The correlation between the values of TL and OL and those of CRL emphasized that the systematic OL measurement in limb long bones correlated better than TL with development age, since OL increased faster than TL. TL and OL also correlated with the CRL by bivariate allometry (ln y = ln a + b ln x) and the data obtained showed that they grew with positive allometry. The comparison between the cumulative values of the bones examined in each limb showed that both TL and OL grew relatively faster in the lower limb than the upper; the greatest growth rate was found for OL in the lower limb. These results many provide a tool for a comprehensive assessment of long bone growth patterns and may be useful in determining fetal growth even in incomplete specimens, in which one or some long bones can still be measured.

Embryonic and Fetal Development↗

Scaling of the mammalian middle ear.

This study considers the general question how animal size limits the size and information receiving capacity of sense organs. To clarify this in the case of the mammalian middle ear, I studied 63 mammalian species, ranging from a small bat to the Indian elephant. I determined the skull mass and the masses of the ossicles malleus, incus and stapes (M, I and S), and measured the tympanic membrane area, A1. The ossicular mass (in mg) is generally negatively allometric to skull mass (in g), the regression equation for the whole material (excluding true seals) being y = 1.373 x(0.513). However, for very small mammals the allometry approaches isometry. Within a group of large mammals no distinct allometry can be discerned. The true seals (Phocidae) are exceptional by having massive ossicles. The size relations within the middle ear are generally rather constant. However, the I/M relation is slightly positively allometric, y = 0.554 x(1.162). Two particularly isometric relations were found; the S/(M + I) relation for the ossicles characterized by the regression equation y = 0.054 x(0.993), and the relation between a two-dimensional measure of the ossicles and the tympanic membrane ares, (M + I)2/3 /A1. As in isometric ears the sound energy collected by the tympanic membrane is linearly related to its area, the latter isometry suggests that, regardless of animal size, a given ossicular cross-sectional area is exposed to a similar sound-induced stress. Possible morphological middle ear adaptations to particular acoustic environments are discussed.

Animals↗

Development of maximum metabolic rate and pulmonary diffusing capacity in the superprecocial Australian Brush Turkey Alectura lathami: an allometric and morphometric study.

The Australian Brush Turkey Alectura lathami is a member of the Megapodiidae, the mound-building birds that produce totally independent, "superprecocial" hatchlings. This study examined the post-hatching development of resting and maximal metabolic rates, and the morphometrically determined changes in pulmonary gas exchange anatomy, in chicks during 3.7 months of growth from hatchlings (122 g) to subadults (1.1 kg). Allometric equations of the form y=aM(b) related gas exchange variables (y) to body mass (M, g). Metabolic rates were measured with open-flow respirometry (mL O2 min(-1)) of chicks resting in the dark and running above the aerobic limit on a treadmill. Resting metabolic rate (RMR=0.02 M(0.99)) and maximal metabolic rate (MMR=0.05 M(1.07)) scaled with exponents significantly above those of interspecific allometries of adult birds. However MMR was below that expected for other species of adult birds in flapping flight, consistent with the Brush Turkey's ground-dwelling habits. Total lung volumes (mL) increased faster than isometrically (V(L)=0.0075 M(1.19)), as did the surface area (cm(2)) of the blood-gas barrier (S(t)=7.80 M(1.23)), but the data overlapped those of adult species. Harmonic mean thickness of the blood-gas barrier was independent of body size (mean tau(ht),=0.39 microm) and was about twice that expected for flying birds. Diffusing capacity (mL O2 min(-1) kPa(-1)) of the blood-gas tissue barrier increased faster than isometrically (Dto2=0.049 M(1.23)); in hatchling Brush Turkeys, it was about 30% expected for adult birds, but this difference disappeared when they became subadults. When compared to altricial Australian pelicans that hatch at similar body masses, superprecocial Brush Turkeys had higher MMR and higher Dto2 at the same body size. A parallel allometry between MMR and Dto2 in Brush Turkeys and pelicans is consistent with the concept of symmorphosis during development.

Animals↗

Facial heights: evolutionary relevance of postnatal ontogeny for facial orientation and skull morphology in humans and chimpanzees.

Facial heights, i.e. the vertical distances between the superior and inferior limits of facial compartments, contribute to the orientation of the viscerocranium in the primate skull. In humans, vertical facial variation is among the main sources of diversity and frequently associated with an integrated suite of other cranio-mandibular traits. Facial heights and kyphosis are also important factors in interspecific variation and models of hominoid evolution. The ontogenetic determination of adult facial orientation and its relation to phylogenetic variation are unclear, but crucial in all previously mentioned respects. We addressed these issues in a sample of 175 humans and chimpanzees with Procrustes based geometric morphometrics, testing hypotheses of interspecific similarity in postnatal ontogenetic trajectories, early versus later ontogenetic facial pattern determination, and a developmental model of morphological integration. We analyzed the contribution of postnatal morphogenesis to adult vertical facial variation by partitioning morphological variation into a portion of pure growth allometry and a non-allometric fraction. A statistically significant difference of growth-allometries revealed that in both species growth established the adult skull proportions by vertical facial expansion, but while in chimpanzees the complete viscerocranium showed reorientation, in humans only the lower face was modified. In both species the results support a hypothesis of early facial pattern determination. A coincident emergence of morphological traits favors a hypothesis of developmental integration of the face, excluding traits of the basi- and neurocranium. Interspecific differences in integration may have implications for evolutionary studies. The present findings indicate that growth establishes the adult skull proportions and integrates principal facial orientation patterns, already there in early postnatal ontogeny.

Adult↗

Metabolic adaptation to hypoxia: cost and benefit of being small.

Following metabolic size allometry, the specific metabolic rate of mammals increases with decreasing body mass, resulting in a steeper metabolic fall-off and a faster exhaustion of energy reserves under hypoxic conditions. However, both mammalian hibernators and fetuses are able to temporarily "switch-off" Kleiber's rule as an adaptation to limited food or oxygen supply. Further exceptions to the usual metabolic size relationship are observed in newborn mammals. For instance, neonatal mouse hearts exhibit slower calorimetric "dying curves" under conditions of ischemia, although their aerobic tissue metabolic rates are higher than in adult samples. This is apparently due to a transient reduction of metabolic rate back to the former feto-maternal level. A continuing deviation from metabolic size allometry is found in newborn marsupials (Monodelphis domestica) where the "inappropriately" low specific metabolic rate is a precondition of efficient growth and tissue aerobiosis in spite of extreme immaturity. Obviously, adaptive suppression of elevated metabolism in organisms of small size results in a dramatic improvement of oxygen supply. Vice-versa, the overall increase in specific metabolic rate with decreasing body size might be regarded as one of several phylogenetic adaptations to protect tissues from hyperoxygenation.

Acclimatization↗

Scaling of bite force in the blacktip shark Carcharhinus limbatus.

Although bite force is a frequently studied performance measure of feeding ecology, changes in bite force over ontogeny have rarely been investigated. Biting by the blacktip shark Carcharhinus limbatus was theoretically modeled over ontogeny to investigate the scaling of bite force, the morphological basis of the observed scaling relationship, the ecological consequences of ontogenetic changes in performance, and whether cranial morphometrics can be used as an accurate proxy for bite force. Theoretical bite force, which was positively allometric with respect to total length (TL), ranged from 32 N (61 cm TL) to 423 N (152 cm TL) at the anterior tips of the jaws and from 107 (61 cm TL) to 1083 N (152 cm TL) at the posterior teeth. This observation is attributed to positive allometry in the mechanical advantage of the jaw-adducting mechanism and the cross-sectional area of all four jaw-adducting muscles. Theoretical bite force was accurately predicted by cranial morphometrics including prebranchial length and head width as well. Although positive allometry of bite force in C. limbatus would seem to indicate an ecological necessity for this phenomenon, dietary analyses do not necessarily indicate any ontogenetic shift in prey types requiring larger bite forces. The positively allometric increase in theoretical bite force may be associated with numerous other selective pressures including maintenance of an apical position within the ecosystem.

Age Factors↗

Growth of capillaries and myocardial cells in the normal rat heart.

The normal growth process of myocardial cells and capillaries in the late postnatal period (after weaning) was studied in 33 normal male Wistar rats (group I: 5 weeks, n = 9; group II: 7 weeks, n = 7; group III: 13 weeks, n = 8; group IV: 52 weeks, n = 9). The rats were fixed by retrograde vascular perfusion via the abdominal aorta. Two transverse and 2 longitudinal sections per animal were selected at random from the left ventricular papillary muscles for light and electron microscopic stereological investigation. Length density and surface density of myocardial cells and capillaries were estimated with correction for partial anisotropy and curvature by means of the mathematical model of a Dimroth-Watson orientation distribution. The results were analyzed by allometric techniques. Relative left and right ventricular weight decreased continuously throughout the growth process; in terms of allometry, ventricular weight was proportional to (body weight). The capillaries showed predominantly longitudinal growth with nearly constant cross-sectional area, whereas the myocardial cells grew by nearly harmonic enlargement into all three directions of space. The relation between total capillary length and left ventricular weight could be described in close approximation by the allometric model (r = 0.98, P less than 0.001). Corresponding to the result that total capillary length was proportional to (left ventricular weight)0.71, the myocardial capillarization (length, surface area, and volume of capillaries per unit tissue volume) decreased with increasing heart size. As allometry makes possible the quantitation of the intensity of growth processes by a single dimensionless number (the allometric exponent), this method permits a convenient interlaboratory comparison of growth studies, and it allows a comparison of capillary reactions during maturation and in experimental cardiac hypertrophy.

Animals↗

Relative growth of the skull and postcranium in giant transgenic mice.

Cross-sectional allometric growth patterns of the cranial and postcranial skeleton were compared between giant transgenic (MT-rGH) mice and their normal littermate controls. Body weights, external body dimensions, and a series of cranial and postcranial linear dimensions of the skeleton were determined for samples of known age. Comparative bivariate and multivariate allometric analyses were completed in order to determine whether (1) the larger transgenic mice differed significantly from the normal controls in aspects of body and skeletal proportions, and (2) any such proportion differences resulted from general allometric effects of overall weight or skeletal size increase. Results demonstrate that the transgenic mice do exhibit significantly different body and skeletal proportions than normal control adults. Allometric comparisons of the skeletal dimensions relative to body weight reveal similar coefficients of growth allometry but several differences in gamma-intercept values in the transgenic vs. control groups. The comparisons among the skeletal dimensions of the skull and postcranium generally reveal the sharing and differential extension of common growth allometries in the two groups. Thus, the elevated levels of growth hormone (GH) and insulin-like growth factor I (IGF-I) in the transgenic mice appear to result in increased overall growth for the various skeletal elements, but in the relative proportions determined by intrinsic growth controls within that system.

Animals↗

An examination of cetacean brain structure with a novel hypothesis correlating thermogenesis to the evolution of a big brain.

This review examines aspects of cetacean brain structure related to behaviour and evolution. Major considerations include cetacean brain-body allometry, structure of the cerebral cortex, the hippocampal formation, specialisations of the cetacean brain related to vocalisations and sleep phenomenology, paleoneurology, and brain-body allometry during cetacean evolution. These data are assimilated to demonstrate that there is no neural basis for the often-asserted high intellectual abilities of cetaceans. Despite this, the cetaceans do have volumetrically large brains. A novel hypothesis regarding the evolution of large brain size in cetaceans is put forward. It is shown that a combination of an unusually high number of glial cells and unihemispheric sleep phenomenology make the cetacean brain an efficient thermogenetic organ, which is needed to counteract heat loss to the water. It is demonstrated that water temperature is the major selection pressure driving an altered scaling of brain and body size and an increased actual brain size in cetaceans. A point in the evolutionary history of cetaceans is identified as the moment in which water temperature became a significant selection pressure in cetacean brain evolution. This occurred at the Archaeoceti - modern cetacean faunal transition. The size, structure and scaling of the cetacean brain continues to be shaped by water temperature in extant cetaceans. The alterations in cetacean brain structure, function and scaling, combined with the imperative of producing offspring that can withstand the rate of heat loss experienced in water, within the genetic confines of eutherian mammal reproductive constraints, provides an explanation for the evolution of the large size of the cetacean brain. These observations provide an alternative to the widely held belief of a correlation between brain size and intelligence in cetaceans.

Animals↗

Elucidation of human amphotericin B pharmacokinetics: identification of a new potential factor affecting interspecies pharmacokinetic scaling.

PURPOSE: To elucidate the pharmacokinetics of amphotericin B in rats, mice and humans, and to perform interspecies scaling to humans using allometry. METHODS: Plasma concentrations following intravenous bolus administration in rats, and mice were determined by HPLC. Human pharmacokinetic parameters elucidated from literature data were validated in a preliminary study involving a patient receiving daily infusion dose for 27 days. A critical literature review was conducted to identify appropriate pharmacokinetic parameter values in other species for interspecies scale-up. Interspecies allometric scale-up was performed across mice, rats, rabbits and dogs and the resulting predictions in humans were compared to observed values. RESULTS: A triexponential decline in rat, mouse and human plasma concentrations were observed. No gender differences in rat pharmacokinetics were observed. In contrast to allometry, mouse CL was smaller (82 vs 116 ml/h/kg) and T0.5 (33 vs 20 h) was longer compared to rat. In the preliminary human study, Cpeak and Cmin values remained relatively constant over the duration of therapy, and a CL, MRT, T0.5, Vss and Vdarea of 26 ml/h/kg, 10 and 23 days, 6.2 and 20 L/kg, respectively, were estimated. The relative contributions of the terminal phase area in rat, mouse and human were 75%, 92% and 31%, respectively. Interspecies allometric scale-up predictions of human CL (41 ml/h/kg), CLu (467 ml/h/kg) and Vss (3.3 L/kg) were similar to reported values, whereas poor predictions of human Vuss (33 L/kg), Vdarea (4.1 L/kg) and T0.5 (3 days) were obtained. CONCLUSIONS: Insignificant accumulation in humans inspite of the long terminal T0.5 was rationalized to be due to the small terminal-phase area contribution. While human CL and Vss were successfully predicted in the interspecies scaling, poor predictions of human Vdarea and T0.5 were obtained, which was attributed to disposition pattern differences between humans and other species, a potential new critical factor affecting interspecies scale-up.

Amphotericin B↗

Scaling of elastic strain energy in kangaroos and the benefits of being big.

Large kangaroos are unique among mammals in their ability to uncouple aerobic metabolic energy costs from the speed of locomotion, making hopping an economical gait. During the first half of the ground-contact phase, kinetic energy lost from the body is stored as elastic strain energy, predominantly in the hind limbs. The subsequent recoil returns kinetic and potential energy to the body. Here we show that the allometry of structures in the legs and feet of Macropodoidea is different from that of quadrupedal eutherian mammals. The potential for elastic energy storage in hoppers is shown to scale with strong positive allometry. This is a function of the structural properties of muscle-tendon units in the distal hind limbs and the postures adopted by hopping kangaroos. Our findings demonstrate how the use of tissue elasticity is strongly mass dependent and help explain the observed energetic phenomena.

Animals↗

Allometric cascade as a unifying principle of body mass effects on metabolism.

The power function of basal metabolic rate scaling is expressed as aM(b), where a corresponds to a scaling constant (intercept), M is body mass, and b is the scaling exponent. The 3/4 power law (the best-fit b value for mammals) was developed from Kleiber's original analysis and, since then, most workers have searched for a single cause to explain the observed allometry. Here we present a multiple-causes model of allometry, where the exponent b is the sum of the influences of multiple contributors to metabolism and control. The relative strength of each contributor, with its own characteristic exponent value, is determined by the control contribution. To illustrate its use, we apply this model to maximum versus basal metabolic rates to explain the differing scaling behaviour of these two biological states in mammals. The main difference in scaling is that, for the basal metabolic rate, the O(2) delivery steps contribute almost nothing to the global b scaling exponent, whereas for the maximum metabolic rate, the O(2) delivery steps significantly increase the global b value.

Adenosine Triphosphate↗