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Size correction: comparing morphological traits among populations and environments.

Morphological relationships change with overall body size and body size often varies among populations. Therefore, quantitative analyses of individual traits from organisms in different populations or environments (e.g., in studies of phenotypic plasticity) often adjust for differences in body size to isolate changes in allometry. Most studies of among population variation in morphology either (1) use analysis of covariance (ANCOVA) with a univariate measure of body size as the covariate, or (2) compare residuals from ordinary least squares regression of each trait against body size or the first principal component of the pooled data (shearing). However, both approaches are problematic. ANCOVA depends on assumptions (small variance in the covariate) that are frequently violated in this context. Residuals analysis assumes that scaling relationships within groups are equal, but this assumption is rarely tested. Furthermore, scaling relationships obtained from pooled data typically mischaracterize within-group scaling relationships. We discuss potential biases imposed by the application of ANCOVA and residuals analysis for quantifying morphological differences, and elaborate and demonstrate a more effective alternative: common principal components analysis combined with Burnaby's back-projection method.

Biological Evolution↗

Variation in enamel thickness and cusp area within human maxillary molars and its bearing on scaling techniques used for studies of enamel thickness between species.

Thirty-nine unworn maxillary molars (M1 = 18, M2 = 12, M3 = 9), representing 37 individuals of a Slavic population, were sectioned through the mesial cusps in a plane perpendicular to the cervical margin of the crown. Whether the range of variation in enamel cross-sectional area between individuals corresponds to the range of variation found between teeth within a species was investigated. Total tooth crown area, as determined by planimetry, was used as a scaling factor on the basis that a biologically meaningful scaling procedure should take into account both enamel thickness and total tooth crown area, rather than the size of the dentine-enamel junction. Uni- and multivariate statistics revealed that the enamel cross-sectional area of second and third molars is absolutely and relatively larger than it is in first upper molars and that it scales close to isometry with tooth crown area. Conversely, first maxillary molars, with their relatively and absolutely thinner enamel, exhibit positive allometry between enamel and tooth crown area. These patterns were retained even when differences in overall tooth size were adjusted for. Hence, the notion that there exists a 'species-specific' amount of enamel over the tooth crown must be reconsidered, and one should refrain from pooling different tooth types for taxonomic purposes. At least two scaling factors are needed to describe human maxillary molars alone. However, this increase in enamel cross-sectional area from anterior to posterior is not unexpected if one bears in mind that posterior teeth occupy a more advantageous position relative to the chewing muscles.(ABSTRACT TRUNCATED AT 250 WORDS)

Austria↗

Mathematical representation of organ growth in the human embryo/fetus.

During human pregnancy, there is a huge increase in the total weight of the embryo/fetus from conception to term. The total growth, which is the summation of growth of the various organs and tissues that make up the organism, was analyzed in a previous paper and fitted to the Gompertz equation for growth. In the present study, allometry, the quantitative representation of the consequence of size, was utilized to describe the correlation of individual fetal organ/tissue weights with the total fetal weight. The organ/tissue weight and the total fetal weight data used in the analyses were pooled from various sources that provided data ranging from 25 days to 300 days post-conception. Allometric equations are presented for 16 embryo/fetal organs and tissues. The standard allometric equation gave adequate fits for embryo/fetal adrenal, bone, bone marrow, brain, heart, liver, pancreas, plasma, skeletal muscle, extracellular water, thymus and thyroid; but it was necessary to use a quadratic form of the allometric equation for embryo/fetal fat, kidney, lung and spleen. Parameters were also calculated for crown-to-rump and crown-to-heels for fetal lengths that occur during pregnancy.

Adipose Tissue↗

A synergic approach to plant pattern generation.

The paper shows convergences between the results found in various models of phyllotaxis. It shows that a synergic approach is needed to deal with the problems of phyllotaxis. An algorithm, called the phi-model, based on the observation of the meaningful and symmetry-generating presence of the golden ratio phi in all types of spiral patterns, and consequently in all types of regular patterns in phyllotaxis, is proposed. The model is suggested by a property of the allometry-type model for pattern recognition in phyllotaxis. It extends recent morphological models developed around the idea of packing efficiency of plant primordia, models that yield the noble numbers, among which are the divergence angles of spiral patterns. The phi-model also gives the noble numbers and moreover orders them in a way that establishes connections with the morphogenetic principles used in models for pattern generation; the order has to do with the relative frequencies of the spiral patterns in nature. The phi-model is a link between the two entropy models in phyllotaxis and offers a nice correspondence with the minimal entropy model generated by a systemic and holistic approach. This latter type of approach is put forward as being able to give a general framework in which to organize the concepts, results, and models in phyllotaxis in a way that produces a synergy of efforts. The necessity of doing so is seen clearly when one considers that phyllotaxis-like patterns are encountered in other fields of research, so that the problem appears to transcend the strict botanical substratum.

Algorithms↗

Some effects of ovariectomy and estrogen replacement on body composition in the rat.

Sprague-Dawley rats were ovariectomized (OvX) at 3 ages, day 2 (D2), week 4 (W4) and week 7 (W7); a group of OvX W7 rats were treated daily with estrogen (OB;2 micrograms for 2 or 5 weeks from 10 weeks of age). Rats were slaughtered at 4 ages, weeks 7, 9, 12 and 15, for the chemical analysis of carcass and skin. Chemical compositions were analysed as % wet weight and as component weights by two-way analysis of variance. Component weights were also analysed by allometry, regressing against nose-anal length. Ovariectomy increased overall body weight without causing obesity. The weight gain of the OvX rat was mainly a true growth response but OvX affected body proportions so that at a given body length the OvX rat had a larger skin and carcass than controls. Ovariectomy at the earliest age (D2) produced the smallest response in body weight and body length but produced the greatest fat redistribution towards the skin and away from the carcass; there was no net change in whole body fat levels following OvX. Long-term daily OB treatment increased fat reserves but slowed the growth of other body components, including the axial skeleton. Whereas OvX redistributed components between skin and carcass, OB treatment reversed this process.

Aging↗

Heart and lung adaptations to pregnancy and lactation in a crocidurine shrew.

Heart and lung mass, rate of oxygen consumption (VO2), respiration rate (fR), tidal volume (VT), and heart rate (fH), were measured at rest and thermoneutrality in the shrew Crocidura russula monacha [(This shrew is claimed to be Crocidura suaveolens (Catzeflis, F., T. Maddalena, S. Hellwing and P. Vogel (1985). Unexpected findings on the taxonomic status of East Mediterranean Crocidura russula auct. (Mammalia, Insectivora). Zeitschrift fur Saugetierkunde 50: 185-201)] in nullipar (N), pregnant (P) and lactating (L) females. The heart mass of N females is large (0.86% of body mass) but fH is slow (70% of that expected by mammalian allometry), while lung mass is small (87% of expected) and fR is high (47% above expected). The 31% higher than expected expired ventilation (VE) matches the 25% higher than expected VO2. In P females the fH of 560 min-1 did not change but it increased in L females to 620 min-1. VE did not change in P females but decreased in L females from 11.3 ml center dot min-1 to 10.1 ml min-1. The normal VT decreased in P and L females from 47.3 mu l to 38.4 mu l and 37.8 mu l respectively. The lower than expected resting fH of N females may provide sufficient scope for increased heart work and oxygen supply during exercise. The large heart may be more efficient. Hyperventilation in N females is indicated by the calculated relatively high fractional concentration of oxygen in expired air (FECO2), and low fractional concentration of CO2 in expired air (FECO2) (17.7% and 3.2% respectively), the high oxygen partial pressure in alveolar gas (PAO2) and low alveolar CO2 (PACO2), 119 and 34 Torr respectively, which facilitate O2 transport through the lung air-blood barrier. The elevated VO2 in P and L females is achieved by increased respiratory efficiency from the normal 15%, to 24% and 29% respectively.

Adaptation, Physiological↗

A return to time, cells, systems, and aging: III. Gompertzian models of biological aging and some possible roles for critical elements.

In this paper, I continue my investigation into the modeling of senescence in biological hierarchies. Making use of my previous discussion on non-reestablishable biological components, I derive a mathematical model which has Gompertzian-like dynamics. I show how this model may be approximated, in certain instances, by a Gompertzian equation. I then demonstrate how our approach yields a biological interpretation for the parameters in the Gompertzian equation. I then demonstrate how changes in the parameter values may be interpreted in light of the biology. Subsequently, I review the literature on the allometry of aging, and I demonstrate how my reliability model may be used to obtain--in a qualitative manner--some of the lifespan curves found in the literature. I close my discussion by constructing a more complex reliability model which incorporates the deterministic failure of biological components with stochastic aspects of senescence.

Aging↗

Quantitative determination by ELISA of tobacco necrosis virus from necrotic local lesions in tobacco.

The amounts of tobacco necrosis virus antigen from necrotic lesions in tobacco leaves were estimated by ELISA. Less than 10% variation among different plates was obtained for the same sample placed in 3 wells randomly selected among the 60 internal wells of the plates, including in each test a dilution series of purified virus. Huxley's simple allometry equations y = bxm (y, absorbance; x, virus concentration) were calculated for each plate and gave reproducible results within a large range of virus concentration. The amount of viral antigen recovered from the necrotic centres of lesions was always significantly lower than those from the living tissues of the halo surrounding the centre. The serological activity recovered from both necrotic and halo tissues was not increased upon various treatments with disaggregating agents. During the continuous growth of the lesions the amounts of viral antigen extracted from the necrotic centre linearly decreased with time, suggesting virus degradation, whereas the amounts of antigen extracted from the living halo tissues increased with time, indicating that mechanisms of restricting viral spread and multiplication were not operating.

Antigens, Viral↗

Water balance and kidney function in the least shrew (Cryptotis parva).

We assessed renal function in least shrews (Cryptotis parva, body mass 4.7 g) within the context of overall water balance. The glomerular filtration rate (GFR) of shrews with unlimited food and water was 2.4 ml/h, about 60% of the rate predicted from body mass. Of this, about 3% (0.075 ml/h) was excreted as urine with an osmolality of 1944 mmol/kg, 5.5 times plasma osmolality. Shrews had a total water turnover (5 ml/day) two to three times higher than expected from allometry for a small mammal of this size. Water influx was partitioned among preformed water from food (65%), drinking (16%), and metabolic water (20%). Water efflux was divided among urine flow (35%), fecal water loss (estimated as 23%), and evaporation (by difference, 42%). Least shrews had a high water turnover rate and relatively high urine flow rate (UFR); this likely reflects a combination of factors, including high metabolism, active lifestyle, and wet diet.

Animal Feed↗

Developing animals flout prominent assumptions of ecological physiology.

Every field of biology has its assumptions, but when they grow to be dogma, they can become constraining. This essay presents data-based challenges to several prominent assumptions of developmental physiologists. The ubiquity of allometry is such an assumption, yet animal development is characterized by rate changes that are counter to allometric predictions. Physiological complexity is assumed to increase with development, but examples are provided showing that complexity can be greatest at intermediate developmental stages. It is assumed that organs have functional equivalency in embryos and adults, yet embryonic structures can have quite different functions than inferred from adults. Another assumption challenged is the duality of neural control (typically sympathetic and parasympathetic), since one of these two regulatory mechanisms typically considerably precedes in development the appearance of the other. A final assumption challenged is the notion that divergent phylogeny creates divergent physiologies in embryos just as in adults, when in fact early in development disparate vertebrate taxa show great quantitative as well as qualitative similarity. Collectively, the inappropriateness of these prominent assumptions based on adult studies suggests that investigation of embryos, larvae and fetuses be conducted with appreciation for their potentially unique physiologies.

Anatomy, Comparative↗

A comparative analysis of temporomandibular joint morphology in the African apes.

A number of researchers have suggested a functional relationship between dietary variation and temporomandibular joint (TMJ) morphology, yet few studies have evaluated TMJ form in the African apes. In this study, I compare TMJ morphology in adults and during ontogeny in Gorilla (G.g. beringei, G.g. graueri, and G.g. gorilla) and Pan (P. paniscus, P. troglodytes troglodytes, P.t. schweinfurthii, and P.t. verus). I test two hypotheses: first, compared to all other African apes, G.g. beringei exhibits TMJ morphologies that would be predicted for a primate that consumes a diet comprised primarily of moderately to very tough, leafy vegetation; and second, all gorillas exhibit the same predicted morphologies compared to Pan. Compared to all adult African apes, G.g. beringei has higher rami and condyles positioned further above the occlusal plane of the mandible, relative to jaw length. Thus, mountain gorillas have the potential to generate relatively more muscle force, more evenly distribute occlusal forces along the postcanine teeth, and generate relatively greater jaw adductor moment. G.g. beringei also exhibits relatively wider mandibular condyles, suggesting these folivorous apes are able to resist relatively greater compressive loads along the lateral and/or medial aspect of the condyle. All gorillas likewise exhibit these same shape differences compared to Pan. These morphological responses are the predicted consequences of intensification of folivory and, as such, provide support for functional hypotheses linking these TMJ morphologies to degree of folivory. The African apes to not, however, demonstrate a systematic pattern of divergence in relative condylar area as a function of intensification of folivory. The ontogenetic trajectories for gorillas are significantly elevated above those of Pan, and to a lesser but still significant degree, mountain gorillas similarly deviate from lowland gorillas (G.g. gorilla and G.g. graueri). Thus, adult shape differences in ramal and condylar heights do not result from the simple extrapolation of common growth allometries relative to jaw length. As such, they are suggestive of an adaptive shift towards a tougher, more folivorous diet. However, the allometric patterning for condylar area and condylar width does not systematically conform to predictions based on dietary specialization. Thus, while differences in condylar shapes may confer functional advantages both during growth and as adults, there is no evidence to suggest selection for altered condylar proportions, independent of the effects of changes in jaw size.

Africa↗

Comparison of hind limb muscle mass in neonate and adult prosimian primates.

Little ontogenetic data exist to indicate whether muscular organization of neonates reflects adult locomotion (e.g., leaping) or infant activities like clinging or the initial quadrupedal phase of locomotion that typifies most infant primates. In the present study, five species of primates with contrasting modes of locomotion were examined. Twenty-eight preserved neonatal and adult cadavers were studied by careful dissection of the hip, thigh, and leg muscles. Wet weights were taken of limb muscles after removal, and the muscles were combined into major functional groups (e.g., flexors, extensors) of each limb segment. Results demonstrate that the distribution of muscle mass within the thigh and within the leg are similar between neonates and adults for all species, with major groups varying by 5% or less in all but two age comparisons. Crural indices of the neonates are nearly identical to those of the adults, but leg/thigh muscle mass ratios were higher in the neonates. Species vary greatly in the percentage of adult limb segment muscle mass present in neonates, with Tarsius syrichta having the greatest percentage for all segments and two lemurids showing the least. These results primarily track differences in relative body mass at birth rather than developmental differences. The adaptive distribution of muscle, as discussed previously for adult prosimians, appears to be established at birth. Neonates of leaping species already have much larger quadriceps muscles than quadrupeds. Differences between large- and small-bodied leapers (e.g., pronounced superficial plantarflexor masses in tarsiers and pronounced deep plantarflexor masses in sifakas) also are present in neonates. Ratios of muscle mass over body mass are smaller in all neonates than in their adult counterparts, suggesting that the neonates are relatively poorly muscled, and that muscle mass must increase with positive allometry during growth.

Aging↗

The eyes of a patrolling butterfly: visual field and eye structure in the Orange Sulphur, Colias eurytheme (Lepidoptera, Pieridae).

Sensory information plays a critical role in determining an animal's behavior on both proximate and evolutionary timescales. Butterflies, like many other insects, use vision extensively over their lifetimes, and yet relatively little work has been published to date on their visual capabilities. We describe the visual system of a pierid butterfly, Colias eurytheme, with the ultimate goal of better understanding its role in shaping the behavior of this animal. We made several measurements: visual field dimensions, eye surface area, interommatidial angle (Deltaphi), facet diameter (D), and eye parameter (p). C. eurytheme had a large visual field and considerable regional variation in visual acuity, as inferred by Deltaphi and D. When compared to females, males had larger eye surface areas, smaller Deltaphi, and larger D in all regions except ventrally. Both sexes had proportionally large eye surface areas compared to other butterflies. Minimum p in males was small, indicating that some regions of their eyes may operate close to the diffraction limit. Finally, we found that both eye surface area and D scaled positively, but with negative allometry to body size. We discuss the relevance of these visual characteristics to the biology and behavior of C. eurytheme.

Animals↗

Density-dependence as a size-independent regulatory mechanism.

The growth function of populations is central in biomathematics. The main dogma is the existence of density-dependence mechanisms, which can be modelled with distinct functional forms that depend on the size of the population. One important class of regulatory functions is the theta-logistic, which generalizes the logistic equation. Using this model as a motivation, this paper introduces a simple dynamical reformulation that generalizes many growth functions. The reformulation consists of two equations, one for population size, and one for the growth rate. Furthermore, the model shows that although population is density-dependent, the dynamics of the growth rate does not depend either on population size, nor on the carrying capacity. Actually, the growth equation is uncoupled from the population size equation, and the model has only two parameters, a Malthusian parameter rho and a competition coefficient theta. Distinct sign combinations of these parameters reproduce not only the family of theta-logistics, but also the van Bertalanffy, Gompertz and Potential Growth equations, among other possibilities. It is also shown that, except for two critical points, there is a general size-scaling relation that includes those appearing in the most important allometric theories, including the recently proposed Metabolic Theory of Ecology. With this model, several issues of general interest are discussed such as the growth of animal population, extinctions, cell growth and allometry, and the effect of environment over a population.

Animals↗

Development of a methodology for assessing the environmental impact of radioactivity in Northern Marine environments.

The requirement to assess the impacts of radioactivity in the environment explicitly and transparently is now generally accepted by the scientific community. A recently developed methodology for achieving this end for marine ecosystems is presented within this paper. With its clear relationship to an overarching system, the marine impact assessment is built around components of environmental transfer, ecodosimetry and radiobiological effects appraisal relying on the use of "reference organisms". Concentration factors (CFs), dynamic models and, in cases where parameters are missing, allometry have been employed in the consideration of radionuclide transfer. Dose conversion coefficients (DCCs) have been derived for selected flora and fauna using, inter alia, dose attenuation and chord distribution functions. The calculated dose-rates can be contextualised through comparison with dose-rates arising from natural background and chronic dose-rates at which biological effects have been observed in selected "umbrella" endpoints.

Animals↗

Woodpecker cavity aeration: a predictive model.

We studied characteristics of the Syrian woodpecker (Dendrocopos syriacus) cavities in the field and a laboratory model, and rates of gas exchange in the laboratory. Night temperature of occupied cavities is 4.3 degrees C higher than empty ones, representing energy savings of approximately 24%. Oxygen conductance (GNO2) of an empty cavity is 7.1 ml[STPD] (Torr h)(-1), and is affected by winds at velocities up to 0.8 m/s. Day and night body temperatures were 42.0 and 40.1 degrees C, respectively. Steady-state O2 consumption rates (MO2) were 3.49 +/- 0.49 and 2.53 +/- 0.26 ml[STPD] (g h)(-1) during day and night respectively -- higher than predicted by allometry. A mathematical model describing PO2 in a cavity, taking into consideration MO2, GNO2, heat convection and wind speed, from the moment birds inhabit it, was developed. It shows that on the average, one woodpecker staying in its cavity at night does not encounter hypoxic conditions. However, in nest cavities with below the average GNO2, with more inhabitants (e.g. during the breeding season), hypoxia may become a problem.

Animals↗

Unifying evolutionary dynamics: from individual stochastic processes to macroscopic models.

A distinctive signature of living systems is Darwinian evolution, that is, a propensity to generate as well as self-select individual diversity. To capture this essential feature of life while describing the dynamics of populations, mathematical models must be rooted in the microscopic, stochastic description of discrete individuals characterized by one or several adaptive traits and interacting with each other. The simplest models assume asexual reproduction and haploid genetics: an offspring usually inherits the trait values of her progenitor, except when a mutation causes the offspring to take a mutation step to new trait values; selection follows from ecological interactions among individuals. Here we present a rigorous construction of the microscopic population process that captures the probabilistic dynamics over continuous time of birth, mutation, and death, as influenced by the trait values of each individual, and interactions between individuals. A by-product of this formal construction is a general algorithm for efficient numerical simulation of the individual-level model. Once the microscopic process is in place, we derive different macroscopic models of adaptive evolution. These models differ in the renormalization they assume, i.e. in the limits taken, in specific orders, on population size, mutation rate, mutation step, while rescaling time accordingly. The macroscopic models also differ in their mathematical nature: deterministic, in the form of ordinary, integro-, or partial differential equations, or probabilistic, like stochastic partial differential equations or superprocesses. These models include extensions of Kimura's equation (and of its approximation for small mutation effects) to frequency- and density-dependent selection. A novel class of macroscopic models obtains when assuming that individual birth and death occur on a short timescale compared with the timescale of typical population growth. On a timescale of very rare mutations, we establish rigorously the models of "trait substitution sequences" and their approximation known as the "canonical equation of adaptive dynamics". We extend these models to account for mutation bias and random drift between multiple evolutionary attractors. The renormalization approach used in this study also opens promising avenues to study and predict patterns of life-history allometries, thereby bridging individual physiology, genetic variation, and ecological interactions in a common evolutionary framework.

Algorithms↗

Insect appendages and comparative ontogenetics.

It is arguable that the evolutionary and ecological success of insects is due in large part to the versatility of their articulated appendages. Recent advances in our understanding of appendage development in Drosophila melanogaster, as well as functional and expression studies in other insect species have begun to frame the general themes of appendage development in the insects. Here, we review current studies that provide for a comparison of limb developmental mechanisms acting at five levels: (1) the specification of ventral appendage primordia; (2) specification of the limb axes; (3) regulation and interactions of genes expressed in specific domains of the proximal-distal axis, such as Distal-less; (4) the specification of appendage identity; and (5) genetic regulation of appendage allometry.

Animals↗