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A simplified model for bivariate complex allometry.

A three-parameter relationship passing through the origin can be derived from the Gompertz growth curve and provide a simplified but useful description of complex allometry, where variates are not proportional to powers of each other as in simple allometry. Partial linearization is often satisfactory and, after a double logarithmic transformation, the allometric relationship is non-linear only with respect to an exponent of complex allometry, D, which reflects the presence (E # 1) or absence (D = 1) of a curvature in log-log space and provides an approximate test of the hypothesis of simple allometry. Examples are presented concerning growth allometry in white rats, metabolic allometry in placental mammals, and size allometry in the Paddlefish, in the North American Marten and in the Painted Turtle.

Animals

Allometry in primates, with emphasis on scaling and the evolution of the brain.

Allometry should be defined broadly as the study of size and its consequences, not narrowly as the application of power functions to the data of growth. Variation in size may be ontogenetic, static or phyletic. Errors of omission and treatment have plagued the study of allometry in primates. Standard texts often treat brain size as an independent measure, ignoring its allometric relation with body size - on this basis, gracile australopithecines have been accorded the mental status of gorillas. Intrinsic allometries of the brain/body are likewise neglected: many authors cite cerebral folding as evidence of man's mental superiority, but folding is a mechanical correlate of brain size itself. Confusion among types of scaling heads errors of treatment in both historical primacy [Dubois' ontogenetic inferences from interspecific curves] and current frequency. The predicted parameters of brain-body plots differ greatly for ontogenetic, intrapopulational, interspecific and phyletic allometries. I then discuss basic trends in bivariate allometry at the ordinal level for internal organ weights, skeletal dimensions, lifespan and fetal weight. In considering the causes of basic bivariate allometries, I examine the reason for differences among types of scaling in brain-body relationships. The interspecific exponent of 0.66 strongly suggests a relationship to body surfaces, but we have no satisfactory explanation for why this should be so. The tripartite ontogenetic plot is a consequence of patterns in neuronal differentiation. We do not know why intraspecific exponents fall between 0.2 and 0.4; several partial explanations have been offered. Multivariate techniques have transcended the pictorial representation of transformed coordinates and offer new, powerful approaches to total allometric patterns. Allometry is most often used as a 'criterion for subtraction'. In order to assess the nature and purpose of an adaptation, we must be able to identify and isolate the aspect of its form that depends both upon its size and the size of the body within which it resides. Cranial indices and limb lengths are misinterpreted when authors apply no correction for body size. The search for a criterion of subtraction has been most diligently pursued in studies of the brain. Clearly, brain size must be assessed by comparison with a 'standard' animal of the same body size. But how shall size be measured, especially in fossils; and how shall a standard animal be construed. I discuss and criticize three methods recently used: RADINSKY'S foramen magnum criterion; Jerison's minimum convex polygons and cephalization quotients; and the indices of progression in comparison with 'basal' insectivores' of BAUCHOT, Stephan and their colleagues.

Animals

[Intracranial allometry and craniologic changes during domestication of silver foxes].

Skull proportions and pattern of intracranial allometry in farm foxes of bred domesticated populations are compared. Changes in skull measurements and bivariate allometry are observed in domesticated males, which in this respect become closer to domesticated females. The changes in the allometry pattern in the domesticated population points to deep modifications of regulatory ontogenetic processes which underlie the allometric growth. However, morphological changes in domesticated foxes cannot be only accounted for by allometry.

Animals

Human mandibular prenatal growth: bivariate and multivariate growth allometry comparing different mandibular dimensions.

Mandibular growth was studied in 36 human fetuses (both sexes) ranging from 13 to 37 weeks of gestation by bivariate and multivariate analyses (bivariate allometry and principal components analysis, PCA). Several mandibular dimensions were measured and correlated with fetal weight. Considering the different mandibular dimensions in sequence of increasing component weights, PCA agreed with bivariate analysis. No mandibular dimension was considered to increase in isometric relationship. PCA showed the following distances with negative allometry: caput mandibulae-gnathion (both sides), gonion-processus coronoideus (both sides), caput mandibulae-processus coronoideus (both sides) and gonion-gnathion (right side). On the other hand, the following dimensions grew with positive allometry: gonion-gnathion (left side) and symphyseal height (both sides). PCA and bivariate analysis showed higher growth rates for the gonion-processus coronoideus distance and symphyseal height on the right side than on the left. All other mandibular dimensions presented more elevated growth rates on the left than on the right side. During the second and third trimesters of prenatal life the mandibular growth was allometrical; the mandibular body grew with more intensity than the ramus in both length and height. The greatest growth rate was found for the height at the symphysis. The angulus mandibulae presented a negative and slight correlation with the other linear dimensions of the mandible during prenatal life.

Biometry

Size increase and form allometry during evolution of ciliate species in the genera Colpoda and Tillina (Ciliophora: Colpodida).

With the absence of a fossil record to provide direct evidence of evolutionary relationships, many researchers have inferred by circular reasoning that body size increase and form allometry have occurred during the evolution of ciliated protozoa. This study establishes ancestor-descendant relationships among five species of the related genera Colpoda and Tillina using ultrastructural characteristics which are not apparently dependent on size and form. This independent test suggests that phylogenetic size increase and phylogenetic form allometry have occurred during the evolution of these ciliates. Interrelationships of body size increase, increase in number of cortical organelles, and form allometry are discussed in reference to the divergence of these ciliate species.

Animals

Asymptotic growth, egg production and trivariate allometry in Esox masquinongy Mitchill.

Multivariate growth models based on the Lotka-Volterra equations seem more complicated than necessary for a fruitful analysis of relative growth data. A simpler and more practical approach combines multivariate allometry with parabolic or asymptotic growth with respect to time. A modified von Bertalanffy curve, X = A [1-exp (-t/D)]C, is compatible with allometry and yields a satisfactory description of somatic growth and egg production in the female muskellunge (Esox masquinongy). A multiplicative variation model appears adequate and partial linearization can be used. While the muskellunge reaches a large size, the present analysis suggests that its growth pattern is limited (asymptotic, determinate). The allometry exponents inferred from the multivariate asymptotic growth model differ slightly from those obtained from principal component analysis or from a trivariate linear structural relationship, but the latter may reflect physiological age rather than physical time.

Age Factors

Tooth scaling and evolutionary dwarfism: an investigation of allometry in human pygmies.

Gould has predicted that in rapidly dwarfed lineages the postcanine teeth exhibit a different scaling pattern than is the normal interspecific trend. His prediction of strong negative allometry has not been frequently tested in quantitative detail. Here we present results of scaling analyses of the molar teeth in African pygmies compared with other Africans of larger size and in Philippine pygmies compared with Filipinos of larger size. We find a pattern of strong negative allometry of tooth size to skull and body size in both these comparisons. This scaling pattern is explained by recourse to the developmental bases (known or inferred) of dwarfing in these populations. Body size decrease is related to low levels of the growth control substance insulin-like growth factor I (IGF-I), which does not appear to affect the size of the dentition. The implications of such developmental information for our understanding of allometric patterns in general, and dwarfing events in particular, are discussed.

Africa, Central

Dogs large and small: the allometry of energy requirements within a single species.

Dogs are unique among mammals in having a 100-fold range in body weight for nonobese adults. This variation makes the calculation of the power function for metabolic body size and hence the allometry of energy requirements a particularly challenging subject. Several functions have been proposed from W0.68 to W0.88 (W = body weight in kg). In the present study we measured the heat output of 22 dogs representing seven breeds, aged 1-10 y with W from 5.8 to 48.8 kg, using a whole-body calorimeter specifically designed for this purpose. Regression of log energy output against log W gave the equation 678 W0.64 (r = 0.96; P less than 0.001), which is considered to represent resting energy expenditure (REE) as kJ/d. If estimates of the energy cost of activity are added to REE, new equations of 655 W0.69 (low activity) and 643 W0.73 (higher activity) are obtained, depending on the amount of activity included in the calculation. From these results we suggest that the allometry of energy requirements of adult dogs is a function of different exponents for REE and the energy cost of activity. It does not appear to exceed W0.75 and may be nearer to W0.67.

Animal Feed

Growth allometry of the organs in giant transgenic mice.

We have analyzed the absolute and relative (allometric) growth of a series of internal organs in giant transgenic (MTrGH) and littermate control mice to determine the general and organ-specific effects of the altered hormonal environment on growth in these rodents. Comparison of cross-sectional growth allometries of organ weights and external body dimensions between the two samples was based on analyses of covariance. We report significantly increased growth in all of the organs and measurements examined except for the brain. Coefficients of growth allometry differ significantly from isometric values in a number of cases, and thus, the adult transgenic mice exhibit body proportions different from those of the adult controls. Most of these shape differences reflect general allometric size increase, but the liver and spleen of the transgenics undergo special enlargement or growth. These results indicate that the primary effect of elevated GH and IGF-I levels is increased overall growth, but in the relative proportions set by the intrinsic controls of individual organs and body regions.

Aging

Dimensions and allometry of testes, epididymides and spermatozoa in the domestic dog (Canis familiaris).

Dimensions of the testes, epididymides and spermatozoa were measured from a sample of 104, mainly cross-bred, domestic dogs ranging in body mass from 5 to 48 kg. Allometric analyses in relation to body mass indicated isometric changes in testicular mass but epididymal mass and the mass of the cauda epididymidis and sperm numbers in the cauda showed significant positive allometry. Total sperm length, seminiferous tubule diameter and the relative proportion of tissues in the testis showed no significant correlation with body mass, but sperm principal piece length showed a low correlation with body mass. We suggest that the positive allometry of epididymal mass and sperm numbers is related to promiscuous mating and sperm competition in stray dogs.

Animals

Multivariate dental allometry in primates.

Disagreement is current over the question of whether relatively large teeth in some large primates are a natural outcome of growth trends instead of an indication of intrinsic differences. A cross-primate survey of dental scaling relative to skull (and inferred body) size is given in this study, using a principal component technique to measure the multivariate growth relation between two sets of data: dental size and cranial size. Cheek teeth are strongly positively allometric in restriced taxonomic groups, especially in cercopithecoids. Conversely, the allometry drops to an almost linear proportional growth relation when variation in diet is controlled.

Animals

Ecological significance of hypometabolism in nonhuman primates: allometry, adaptation, and deviant diets.

The "Kleiber relationship" describes the interspecific allometry between body size and metabolism. Like other allometric relationships, the Kleiber relationship not only summarizes scaling effects across species but also provides a standard by which species can be compared. One well-noted deviation from the Kleiber relationship is "hypometabolism": metabolic rates below that expected for a given size. It has been suggested in the literature that hypometabolism may be a primitive mammalian trait, a thermoregulatory adaptation, an adaptation to arboreal folivory, or an adaptation to a diet that is deviant for body size. Data on primate physiology and behavior are used to evaluate these hypotheses. Only the deviant-diet hypothesis is supported by the data on nonhuman primates. Indeed, the Jarman-Bell relationship, which is the basis for this hypothesis, provides a more coherent explanation of correlated features of animal physiology and behavior than do the alternative models. Hypometabolism may be an energy-conserving adaptation to a variety of nutritional stresses. The present analysis underscores the point that metabolic rate, like foraging behavior, should be thought of as evolutionarily labile.

Adaptation, Physiological

Allometry of the limb long bones of insectivores and rodents.

In an attempt to investigate the relationships between allometry and locomotory adaptations, we studied the long limb bones of 45 species of insectivores and rodents. Animals ranged from a few grams to about 50 kilograms. Diameter and length of the bones and body mass (when known) were recorded. Regressions of diameter to length, diameter to body mass, and length to body mass were calculated by the least-squares and Model II, or major axis, methods. The results obtained do not agree with the predictions of either the theory of geometric similarity or the theory of elastic similarity. The discrepancies could be due to the fact that animals studied exhibit various modes of locomotion. Moreover, the allometric relationships of the different locomotor patterns are better reflected in insectivores and rodents than in other groups of mammals. The use of a single regression analysis seems to be inadequate when the sample includes a large range of body sizes.

Animals

Allometry of the perivascular spaces of the putamen in aging.

The equation of simple allometry was used to study the relative growths of the diameters of the perivascular spaces (PSs), the diameters of the arterial vessels and their wall thicknesses in specimens of the putamen from 24 subjects aged 62 to 90 years. Our findings supported the view that the diameters of the PSs were mainly related to the dimensions of their arteries, especially in the elderly, and that they grew isometrically with arterial diameters. In addition, the diameters of the PSs were basically independent of age.

Aged

Relative growth of adenomatous polyps of the colon. Stereology and allometry of multiple polyposis.

The volume of the adenomatous mucosa (V), the area of the surface epithelium (Ss), the area of the glandular epithelium (Sg), and the Sg: Ss ratio were calculated in a series of 14 adenomatous polyps (APs) of a case of multiple polyposis of the colon. The equation of simple allometry was used to study the relative growth of the four series of values. Ss grew isometrically with size; Sg overgrew Ss and accounted for most of the increase in V. The Sg:Ss ratio increased with Sg and V.

Colonic Polyps

Basal metabolic rates in mammals: taxonomic differences in the allometry of BMR and body mass.

No single equation adequately describes the allometric relation between body mass and BMR for mammals. Least squares regression of log-transformed data for 248 eutherian species results in a line with a slope (-0.30) significantly different from that of Kleiber's line (-0.25). Interordinal comparisons of least squares regressions of log-transformed BMR and mass suggest that the Insectivora have a significantly steeper slope to their allometric relationship than do most other orders, while the non-insectivore orders are statistically homogeneous with respect to slope. With respect to elevation, Edentata have the lowest BMRs; Marsupialia, Primates and Chiroptera are indistinguishable from each other but above the edentates; Primates, Chiroptera, Rodentia, Lagomorpha and Carnivora form the next highest homogeneous grouping; and Artiodactyla have the highest BMRs, significantly greater than all but Lagomorpha and Carnivora. Analysis of intraordinal variation within the Rodentia suggests significant heterogeneity among families in BMR-mass allometry.

Animals

Allometry of resting metabolic rate in cockroaches.

1. Resting VO2 at 22.7 degrees C was 0.217 +/- 0.007 ml O2/g/hr in Nauphoeta cinerea of 0.51 +/- 0.008 g body mass. 2. Whole animal resting metabolic rate for 11 cockroach species scaled allometrically: VO2 = 0.261 M0.776. 3. Allometry of resting VO2 among cockroach species is similar to that in vertebrates, and is consistent with models based on both "elastic similarity" and "biological similitude."

Animals