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Effects of age and size in the ears of gekkonomorph lizards: middle-ear morphology with evolutionary implications.

The function of the ear depends in part on its absolute size and internal proportions. Thus, in both young individuals and small species, the middle ear is expected to be allometrically enlarged despite its smaller absolute size. Here we aim to compare the ontogenetic allometry of relevant middle-ear structures as observed within gecko (gekkonomorph lizards) species, with the evolutionary allometry observed interspecifically. These observations also provide middle-ear data for future evaluation of variation in auditory sensitivity. The material comprised 84 museum specimens of geckos, representing nine species of three gekkonomorph subfamilies. The results of dissections and measurements show that different reports notwithstanding, the middle-ear ossicular chain is indeed structured as described for geckos by Werner and Wever. Some sexual dimorphism is indicated, but this requires further study. During postnatal ontogeny, the allometric growth in the ratio of the columellar footplate area to body length differed between the intraspecific and interspecific levels, hence species differences in the middle ear do not merely result from animal size. The ratio of the tympanic membrane area to the columellar footplate area increased during ontogeny. In this, geckos resemble birds and probably also mammals. Similarly, when the comparison was among adults representing different species, the ratio of the tympanic membrane area to the columellar footplate area increased with body size. In this, however, the geckos differed from birds and mammals, in which this ratio varied taxonomically, irrespective of body size. It would thus seem that middle-ear proportions have evolved among geckos to produce small interspecific differences, but among amniote tetrapods they have evolved according to different principles in the classes reptiles, birds, and mammals.

Aging↗

Pharmacokinetic scaling of SJ-8029, a novel anticancer agent possessing microtubule and topoisomerase inhibiting activities, by species-invariant time methods.

This study examined the pharmacokinetic disposition of SJ-8029, a novel anticancer agent possessing microtubule and topoisomerase inhibiting activities, in mice, rats, rabbits and dogs after i.v. administration. The serum concentration-time curves of SJ-8029 were best described by tri-exponential equations in all these animal species. The mean Cl, V(ss) and t(1/2) were 0.3 l/h, 0.1 l and 63.2 min in mice, 1.5 l/h, 1.6 l and 247.7 min in rats, 13.8 l/h, 39.6 l and 245.9 min in rabbits, and 29.2 l/h, 44.6 l and 117.4 min in dogs, respectively. Based on animal data, the pharmacokinetics of SJ-8029 were predicted in humans using simple allometry and also by several species-invariant time transformations using kallynochron, apolysichron and dienetichron times. The human pharmacokinetic parameters of Cl, V(ss) and t(1/2) predicted by the simple allometry and various species-invariant time methods were 50.4-145.0 l/h, 369.0-579.8 l and 242.0-1448.3 min, respectively. These preliminary parameter values may be useful in designing early pharmacokinetic studies of SJ-8029 in humans.

Acridines↗

Prenatal sex ratios influence sexual dimorphism in a reptile.

The prenatal environment influences offspring traits in a variety of ways and in a wide range of taxa. For example, maternal allocation of steroids to the eggs influences offspring traits in birds, and in some mammals the intrauterine position influences morphological, behavioural, and physiological traits due to sex-related steroid transfer between sibling fetuses. We show that similar phenomena occur in the common lizard (Lacerta vivipara), a viviparous reptile. Females developing in male-biased clutches had a more masculine allometry (relatively larger heads) at parturition than females developing in female-biased clutches. Males were correspondingly feminized in female-biased clutches. The effects could either be due to diffusion of steroids produced by the offspring or by a general tendency for females to allocate steroids according to the sex ratio of her clutch. Subsequent to parturition, the sexes differed in their growth trajectories depending on sex ratio environment. In males, the difference in allometry between sex ratio environments remained over time, whereas in females the corresponding effect disappeared.

Animals↗

Scaling of body frontal area and body width in birds.

By analyzing a homogenous dataset we show, in contradiction to a previous study, that the scaling of body frontal area (S(b)) with body mass (m(b)) does not differ between passerine and nonpasserine birds. It is likely that comparison of data collected from live passerines with data collected from frozen nonpasserines had led to the incorrect conclusion that the scaling of S(b) varied between the taxa. We suggest that body dimensions collected from frozen specimens, or specimens stored in alcohol, are not applicable to live birds, and that both the current equations presented in the literature for predicting S(b) from m(b) may lead to inaccurate estimates. Using data from preserved specimens, we found that S(b) scales isometrically with m(b) (S(b) proportional, variant m(b) (0.66)), and therefore we found no evidence for larger birds being more streamlined than smaller birds. S(b) scales with negative allometry against wingspan (b), however, and b scales with positive allometry against m(b), so larger birds have smaller S(b) relative to b. In addition, it appears that dorsoventral flattening of the body is a general characteristic of bird's bodies but that it is more pronounced in larger birds, suggesting perhaps a function in terms of increased lift during forward flight. It appears that bird's bodies obey the surface-to-area geometric scaling law, but bird body shape may vary in relation to aerodynamic function. We suggest that a large-scale study, simultaneously measuring S(b) and m(b) in live passerines and nonpasserines, is required to improve the predictive power of S(b) upon m(b) scaling equations, which play a key role in the estimation of mechanical power consumption in flight in birds. Furthermore, the relations between bird body shape and axial skeleton dimensions, with reference to aerodynamic adaptation, warrant further investigation.

Animals↗

Convergent evolution of sexual shape dimorphism in Diptera.

Several patterns of sexual shape dimorphism, such as male body elongation, eye stalks, or extensions of the exoskeleton, have evolved repeatedly in the true flies (Diptera). Although these dimorphisms may have evolved in response to sexual selection on male body shape, conserved genetic factors may have contributed to this convergent evolution, resulting in stronger phenotypic convergence than might be expected from functional requirements alone. I compared phenotypic variation in body shape in two distantly related species exhibiting sexually dimorphic body elongation: Prochyliza xanthostoma (Piophilidae) and Telostylinus angusticollis (Neriidae). Although sexual selection appears to act differently on male body shape in these species, they exhibited strikingly similar patterns of sexual dimorphism. Likewise, patterns of within-sex shape variation were similar in the two species, particularly in males: relative elongation of the male head capsule, antenna, and legs was associated with reduced head capsule width and wing length, but was nearly independent of variation in thorax length. However, the two species presented contrasting patterns of static allometry: male sexual traits exhibited elevated allometric slopes in T. angusticollis, but not in P. xanthostoma. These results suggest that a shared pattern of covariation among traits may have channeled the evolution of sexually dimorphic body elongation in these species. Nonetheless, static allometries may have been shaped by species-specific selection pressures or genetic architectures.

Animals↗

Miniaturization and its effects on cranial morphology in plethodontid salamanders, genus Thorius (Amphibia, Plethodontidae): II. The fate of the brain and sense organs and their role in skull morphogenesis and evolution.

Relative size and arrangement of the brain and paired sense organs are examined in three species of Thorius, a genus of minute, terrestrial salamanders that are among the smallest extant tailed tetrapods. Analogous measurements of representative species of three related genera of larger tropical (Pseudoeurycea, Chiropterotriton) and temperate (Plethodon) salamanders are used to identify changes in gross morphology of the brain and sense organs that have accompanied the evolution of decreased head size in Thorius and their relation to associated changes in skull morphology. In adult Thorius, relative size (area measured in frontal plane, and length) of the eyes, otic capsules, and brain each is greater than in adults of all of the larger genera; relative size of the nasal capsules is unchanged or slightly smaller. Interspecific scaling phenomena--negative allometry of otic capsule, eye and brain size, isometry or slight positive allometry of nasal capsule size, all with respect to skull length--also are characteristic of intraspecific (ontogenetic) comparisons in both T. narisovalis and Pseudoeurycea goebeli. Predominance of the brain and eyes in Thorius results in greater contact and overlap among these structures and the nasal capsules in the anterior portion of the head. This is associated with anterior displacement of both the eyes and nasal capsules, which now protrude anterior to the skull proper; a change in eye shape; and medial deformation of anterior braincase walls. Posteriorly, predominance of the otic capsules has effected a reorientation of the jaw suspensorium to a fully vertical position that is correlated with the novel presence of a posteriorly directed squamosal process and shift in origin of the quadropectoralis muscle. Many of these changes in cranial morphology may be explained simply as results of mechanical (physical) interactions among the skeletal, nervous, and sensory components during head development at reduced size. This provides further evidence of the role of nervous, sensory, and other "soft" tissues in cranial skeletal morphogenesis, and reinforces the need to consider these tissues in analyses of skull evolution.

Animals↗

Interspecies scaling of biliary excreted drugs.

The objective of this study is to predict clearance of drugs in humans from animals which are excreted in the bile. Clearance (CL) of eight drugs known to be excreted in the bile were randomly selected from the literature. Scaling of CL was performed using at least three animal species. Using simple allometry, CL x mean life-span potential (MLP) or CL x brain weight, CLs of studied drugs were predicted in humans. The choice of one of the methods depended on the 'rule of exponents' as described by Mahmood and Balian. A 'correction factor' was calculated by adjusting bile flow rate based on the species body weight (bile flow = mL/day/kg body weight) or liver weight (bile flow = mL/day/kg liver weight). Using the 'rule of exponents' and combining it with the 'correction factor', the CLs of biliary excreted drugs were predicted in humans. Predicted CLs in humans from animals using simple allometry were several times higher for all eight drugs (% error [range] = 46-1703). Using the 'rule of exponents' and combining it with a 'correction factor' as described in this report provided a substantial improvement (% error [range] = 5-91) in the prediction of CL for biliary excreted drugs. The results of this study indicate that the CL of a biliary excreted drug may be overpredicted in humans and by applying the 'correction factor' employed here, the predictability of drug CL in humans from animal data may be significantly improved.

Algorithms↗

A comprehensive analysis of the role of correction factors in the allometric predictivity of clearance from rat, dog, and monkey to humans.

This study was conducted to comprehensively evaluate the performance of various allometric scaling methods for the prediction of human clearance. Allometric scaling was used to predict clearance for 103 compounds, for which clearance data in the rat, dog, monkey, and humans were available. Allometry was performed using all three preclinical species and with combinations of any two species. The methods employed included standard allometry and various correction factors, including brain weight, maximum lifespan potential, and glomerular filtration. Scaling was performed on all compounds universally and on segregated subsets based on allometric exponent, clearance, physicochemical property, or route of elimination. 776 allometric combinations with 27,313 individual outcomes were performed. A predicted-to-observed clearance ratio of 0.5 to twofold was preselected as the criterion for predictive success. The success rate of allometric scaling ranged from 18 to 53%; none of the correction factors resulted in substantially improved predictivity. Furthermore, none of the methods attempted in this study achieved a success rate greater than that observed by simply estimating human clearance based on monkey hepatic extraction. Prospective allometric scaling, with or without correction factors, represents a suboptimal technique for estimating human clearance based on in vivo preclinical data.

Animals↗

Interspecies scaling of biliary excreted drugs: a comparison of several methods.

The objective of this study was to evaluate the predictive performance of several methods of interspecies scaling for the prediction of human clearance of drugs which are excreted in the bile. Ten methods of allometric scaling were used to predict human clearance of biliary excreted drugs from animal data. The methods included the simple allometric approach; the rule of exponents; the rule of exponents with a correction factor; the ratio of human and monkey liver blood flow x monkey clearance; product of clearance and bile flow; product of clearance and UDGPT; product of clearance, bile flow, and UDGPT; and a modified version of the last three approaches in association with the rule of exponents. The results of the study indicate that among these ten approaches, the rule of exponents with the correction factor is the best approach for the prediction of human clearance for drugs which are excreted in the bile. The worst approach is the product of clearance, bile flow, and UDGPT. The simple allometry and the monkey liver blood flow (MLBF) approaches gave almost similar results. For some drugs the simple allometry predicted clearance better than the MLBF approach and vice versa.

Administration, Oral↗

Similarity principles and intrinsic geometries: contrasting approaches to interspecies scaling.

We criticize standard allometric approaches on the grounds that they emphasize scaling to one variable at a time, whereas chemically reactive hydrodynamic systems involved in pharmacokinetic phenomena are of higher dimension. We show that attempts based on mechanical similitude to set a dosage that would be equivalent across species (for example, from mouse to humans) lead to ambiguous results. Another failing of standard allometry may be its incapability to accommodate the neoteny of Homo sapiens, even though it helped discover the phenomenon. The retarded development in our species implied by neoteny can most clearly be seen in the evidence that both our brain size and our lifespan lie well above the allometric curve for Class Mammalia for these features. In contrast to allometry, which proposes a search for scaling coefficients through invariant external measurement reference frames, we propose a search for transformations of coordinate space coefficients in an intrinsic geometry for the mammalian body plan.

Aging↗

Estimation of African ape body length from femur length.

The estimation from long bone lengths of stature in humans or body size in apes has a deep history in physical anthropology. To date, we can enumerate at least five different statistical methods for making such estimations. These methods are: (1) the regression of body length on long bone length (inverse calibration), (2) regression of long bone length on body length followed by solving for body length (classical calibration), (3) major axis regression of body length on long bone length, (4) reduced major axis regression of body length on long bone length, and (5) use of a long bone/body length ratio. We examine some of the statistical properties of these estimators using a large sample of humans (n = 2053) to derive the estimators, and applying them to smaller samples of Pan troglodytes (n = 42), Pan paniscus (n = 8), and Gorilla gorilla (n = 35). Based on the root mean-squared error (RMSE), the reduced major axis is the preferred estimator for body length in the combined Pan sample. However, inverse calibration is the best estimator for body length in gorillas based on the RMSE. Many estimators grossly underestimate body length in the apes. Differences in allometries between humans and great apes are obvious, but it is important to show the assumptions necessary in estimating body size from fossil remains, especially when isolated long bones are recovered and the global allometry is consequently unknown.

Africa↗

A quantitative morphometric comparative analysis of the primate temporal lobe.

Given their importance in language comprehension, the human temporal lobes and/or some of their component structures might be expected to be larger than allometric predictions for a nonhuman anthropoid brain of human size. Whole brain, T1-weighted MRI scans were collected from 44 living anthropoid primates spanning 11 species. Easyvision software (Philips Medical Systems, The Netherlands) was used to measure the volume of the entire brain, the temporal lobes, the superior temporal gyri, and the temporal lobe white matter. The surface areas of both the entire temporal lobe and the superior temporal gyrus were also measured, as was temporal cortical gyrification. Allometric regressions of temporal lobe structures on brain volume consistently showed apes and monkeys to scale along different trajectories, with the monkeys typically lying at a higher elevation than the apes. Within the temporal lobe, overall volume, surface area, and white matter volume were significantly larger in humans than predicted by the ape regression lines. The largest departure from allometry in humans was for the temporal lobe white matter volume which, in addition to being significantly larger than predicted for brain size, was also significantly larger than predicted for temporal lobe volume. Among the nonhuman primate sample, Cebus have small temporal lobes for their brain size, and Macaca and Papio have large superior temporal gyri for their brain size. The observed departures from allometry might reflect neurobiological adaptations supporting species-specific communication in both humans and old world monkeys.

Anatomy, Comparative↗

Patterns of cranial shape variation in the Papionini (Primates: Cercopithecinae).

Traditional classifications of the Old World monkey tribe Papionini (Primates: Cercopithecinae) recognized the mangabey genera Cercocebus and Lophocebus as sister taxa. However, molecular studies have consistently found the mangabeys to be diphyletic, with Cercocebus and Mandrillus forming a clade to the exclusion of all other papionins. Recent studies have identified cranial and postcranial features which distinguish the Cercocebus-Mandrillus clade, however the detailed similarities in cranial shape between the mangabey genera are more difficult to reconcile with the molecular evidence. Given the large size differential between members of the papionin molecular clades, it has frequently been suggested that allometric effects account for homoplasy in papionin cranial form. A combination of geometric morphometric, bivariate, and multivariate methods was used to evaluate the hypothesis that allometric scaling contributes to craniofacial similarities between like-sized papionin taxa. Patterns of allometric and size-independent cranial shape variation were subsequently described and related to known papionin phylogenetic relationships and patterns of development. Results confirm that allometric scaling of craniofacial shape characterized by positive facial allometry and negative neurocranial allometry is present across adult papionins. Pairwise comparisons of regression lines among genera revealed considerable homogeneity of scaling within the Papionini, however statistically significant differences in regression lines also were noted. In particular, Cercocebus and Lophocebus exhibit a shared slope and significant vertical displacement of their allometric lines relative to other papionins. These findings give no support to narrowly construed hypotheses of uniquely shared patterns of allometric scaling, either between sister taxa or across all papionins. However, more general allometric trends do appear to account for a substantial proportion of papionin cranial shape variation, most notably in those features which have influenced traditional morphological phylogenies. Examination of size-uncorrelated shape variation gives no clear support to molecular phylogenies, but underscores the absence of morphometric similarities between the mangabey genera when size effects are controlled. Patterns of allometric and size-uncorrelated shape variation indicate conservatism of cranial form in non- Theropithecus papionins, and suggest that Papio represents the primitive morphometric pattern for the African papionins. Lophocebus exhibits a divergent morphometric pattern, clearly distinguishable from other papionins, most notably Cercocebus. These results clarify patterns of cranial shape variation among the extant Papionini and lay the groundwork for studies of related fossil taxa.

Animals↗

Biological power laws and Darwin's principle.

Assuming that the repertoire of responses by living systems to perturbation gives a measure of their Darwinian fitness in a rapidly fluctuating environment, those that fulfill allometries (power laws) are described by means of catastrophes, whose variables and parameters are smooth functions of biological attributes. Using empirical allometries from a given system as input, a method is proposed to construct its associated catastrophe, allowing specific predictions on its susceptibility to perturbation and related properties, based on general results from catastrophe theory. The method is discussed within the macroecological context, and an example is provided by applying it to ecological systems that satisfy the self-thinning rule.

Animals↗

Sonographic quantitative analysis of the heart in the third trimester of gestation.

The fetal heart was studied with transabdominal sonography in 40 single normal pregnancies to analyze the increase of cardiac length and width (major longitudinal and transverse diameters of the heart) during the third trimester. Relationships between the cardiac data and fetal age, femur length and biparietal diameter (BPD) were explored by allometry and linear regression analysis. The length and width of the heart could be measured reproducibly in the 4-chamber view of the heart. This study verified that the heart grows very rapidly during the third trimester (positive allometry). The linear increase of the fetal heart shows a strong and significant correlation with the increase of both femur length and BPD. This suggests that non invasive analysis of cardiac data can be useful for prenatal detection of congenital heart disease or for the assessment of gestational age.

Female↗

Decapod crustacean chelipeds: an overview.

The structure, growth, differentiation and function of crustacean chelipeds are reviewed. In many decapod crustaceans growth of chelae is isometric with allometry level reaching unity till the puberty moult. Afterwards the same trend continues in females, while in males there is a marked spurt in the level of allometry accompanied by a sudden increase in the relative size of chelae. Subsequently they are differentiated morphologically into crusher and cutter making them heterochelous and sexually dimorphic. Of the two, the major chela is used during agonistic encounters while the minor is used for prey capture and grooming. Various biotic and abiotic factors exert a negative effect on cheliped growth. The dimorphic growth pattern of chelae can be adversely affected by factors such as parasitic infection and substrate conditions. Display patterns of chelipeds have an important role in agonistic and aggressive interactions. Of the five pairs of pereiopods, the chelae are versatile organs of offence and defence which also make them the most vulnerable for autotomy. Regeneration of the autotomized chelipeds imposes an additional energy demand called "regeneration load" on the incumbent, altering energy allocation for somatic and/or reproductive processes. Partial withdrawal of chelae leading to incomplete exuviation is reported for the first time in the laboratory and field in Macrobrachium species.

Agonistic Behavior↗

Driving forces from soil invertebrates to ecosystem functioning: the allometric perspective.

The European soil policy is being focussed towards a more conscious and sustainable use of the soil, taking into account ecological, economical and societal dimensions. Living soil organisms are reliable bioindicators, as they provide the best reflection of the soil system, ecological services and ecosystem functioning therein. These most complex (bio)physical systems indicate, among others, the energy flow. Such processes can be described by rather simple power law relationships. In fact, the average body mass (dry weight) can be seen as an inherent species property, while population density is a much more flexible parameter reflecting ecosystem state. In this study, I review the interactions between these items in relation to feedbacks and conjectured relationships which can be seen as ecological networks. From this novel perspective, allometry can be used as an integrated measure for the anthropogenic influence on landscapes and related food webs. Allometry is, therefore, a perfect surrogate for land use intensity in modelling of field effects for restoration ecology and conservation biology. Robust correlations will be addressed between the density dependence of invertebrates and the ability of soil systems themselves to recover after disturbance. Quantitative indicators of soil community composition and related ecological services are proposed and their application for ecological risk assessment is illustrated.

Animals↗

Hearing dimorphism, trait variation and conflicts over space in the thorax of the bushcricket Requena verticalis (Listroscelidinae: Tettigoniidae: Orthoptera).

The hearing system of Requena verticalis is sexually dimorphic. Previous work has shown size of the auditory spiracle determines absolute threshold and as female spiracles are, on average, larger than males, females are more sensitive to the main energy of the male call. In all measured traits in morphology and physiology, females showed lower coefficient of variation than males. This difference was significant for bulla volume and hearing threshold. In addition, female ear size covaries with thorax dimensions but this is not so in males. Such a finding suggests stabilising selection on ear size in females, perhaps explained by the requirement of females to recognise and locate the male. As the auditory bulla is larger in females than males, so occupying thoracic space, we suggest a possible trade-off in this brachypterous species between hearing sensitivity and sound production. Finally, we examine relative growth of body structures not associated with hearing and those that influence hearing sensitivity. Scaling, where traits are under strong selection, may result in allometry. Female hearing traits show positive allometry with absolute size and while the relationship between bulla volume and spiracle area was positively allometric in females this was not the case for males.

Animal Communication↗