Metabolic rate, sleep duration, and body temperature in evolution of mammals and birds: the influence of geological time of principal groups divergence.

This study contains an analysis of basal metabolic rate (BMR) in 1817 endothermic species. The aim was to establish how metabolic scaling varies between the main groups of endotherms during evolution. The data for all the considered groups were combined and the common exponent in the allometric relationship between the BMR and body weight was established as b = 0.7248. Reduced to the common slope, the relative metabolic rate forms the following series: Neognathae - Passeriformes - 1.00, Neognathae - Non-Passeriformes - 0.75, Palaeognathae - 0.53, Eutheria - 0.57, Marsupialia - 0.44, and Monotremata - 0.26. The main finding is that the metabolic rate in the six main groups of mammals and birds consistently increases as the geological time of the group's divergence approaches the present. In parallel, the average body temperature in the group rises, the duration of sleep decreases and the duration of activity increases. BMR in a taxon correlates with its evolutionary age: the later a clade diverged, the higher is its metabolic rate and the longer is its activity period; group exponents decrease as group divergence nears present times while with increase metabolic rate during activity, they not only do not decrease but can increase. Sleep duration in mammals was on average 40% longer than in birds while BMR, in contrast, was 40% higher in birds. The evolution of metabolic scaling, body temperature, sleep duration, and activity during the development of endothermic life forms is demonstrated, allowing for a better understanding of the underlying principles of endothermy formation.

Metabolic Scaling in Birds and Mammals: How Taxon Divergence Time, Phylogeny, and Metabolic Rate Affect the Relationship between Scaling Exponents and Intercepts.

Analysis of metabolic scaling in currently living endothermic animal species allowed us to show how the relationship between body mass and the basal metabolic rate (BMR) has evolved in the history of endothermic vertebrates. We compared six taxonomic groups according to their energetic characteristics and the time of evolutionary divergence. We transformed the slope of the regression lines to the common value and analyzed three criteria for comparing BMR of different taxa regardless of body size. Correlation between average field metabolic rate (FMR) of the group and its average BMR was shown. We evaluated the efficiency of self-maintenance in ordinary life (defined BMR/FMR) in six main groups of endotherms. Our study has shown that metabolic scaling in the main groups of endothermic animals correlates with their evolutionary age: the younger the group, the higher the metabolic rate, but the rate increases more slowly with increasing body weight. We found negative linear relationship for scaling exponents and the allometric coefficient in five groups of endotherms: in units of mL O2/h per g, in relative units of allometric coefficients, and also in level or scaling elevation. Mammals that diverged from the main vertebrate stem earlier have a higher "b" exponent than later divergent birds. A new approach using three criteria for comparing BMR of different taxa regardless of body mass will be useful for many biological size-scaling relationships that follow the power function.

Evolution of metabolic scaling among the tetrapod: effect of phylogeny, the geologic time of class formation, and uniformity of species within a class.

The metabolic scaling in the animal has been discussed for over 90 years, but no consensus has been reached. Our analysis of 2126 species of vertebrates reveals a significant allometric exponent heterogeneity. We show that classes of terrestrial vertebrates exhibit the evolution of metabolic scaling. Both the allometric coefficient "a" and the allometric exponent "b" change naturally, but differently depending on the geological time of group formation. The allometric coefficient "a" shows the measure of the evolutionary development of systems that forms resting metabolism in animals. Endothermic classes, such as birds and mammals, have a metabolic rate that is in an order of magnitude higher than that in ectothermic classes, including amphibians and reptiles. In the terrestrial vertebrate phylogeny, we find that the metabolic scaling is characterized by 3 main allometric exponent values: b = 3/4 (mammals), b > 3/4 (ectotherms, such as amphibians and reptiles), and b < 3/4 (birds). The heterogeneity of the allometric exponent is a natural phenomenon associated with the general evolution of vertebrates. The scaling factor decreases depending on both the external design and the size (birds vs mammals) of the animal. The metabolic rate and uniformity of species within a class increase as the geological start date of formation of the class approaches the present time. The higher the mass-specific standard metabolic rate in the class, the slower metabolic rate grows with increasing body size in this class. Our results lay the groundwork for further exploration of the evolutionary and ecological aspects of the development of metabolic scaling in animals.

Total Evaporative Water Loss in Birds at Different Ambient Temperatures: Allometric and Stoichiometric Approaches.

Valery M. Gavrilov (2017) Total evaporative water loss (TEWL) in Passeriformes and Non-Passeriformes was estimated by simultaneous measurements of energy expenditure and mass loss in resting birds. It was found that the percentage of heat dissipated by water evaporation depends on body size. Published data for 102 bird species were analyzed together with my own measurements for 157 bird species at thermally neutral temperatures (mostly 25°C) to establish the following relationship between TEWL and body mass: TEWL25°C Aves = 0.28 m0.701, R 2 = 0.92, where TEWL is in g H2O/day and m is body mass (g). The scaling exponent 0.701 ± 0.007 is 0.05 greater than for the relationship of basal metabolic rate (BMR) to body mass. It was found that TEWL in passerines is higher than in non-passerines at all ambient temperatures by 50% at 25°C, 30% at 0°C, 39% at the lower critical temperature, and 59% at the upper critical temperature. The dependence of water loss on body mass at different ambient temperatures (T A) was found to vary in the same manner as evaporative heat loss. TEWL in Passeriformes is approximately 25-60% higher than in Non-Passeriformes (particularly at high T A), which is consistent with the ratio of their BMR levels. Within the thermoneutral zone, the proportion of heat dissipated by evaporation increases by approximately 2.6-fold in small passerines and by almost 4.1-fold in large passerines with the transition from the lower to upper critical temperature. In non-passerines, the proportion of evaporative heat losses increases by approximately 2.7 times within the thermoneutral zone in both large and small birds. The high basal metabolic rate in Passeriformes involves benefits like a higher maximum metabolic power and the ability to breed at lower ambient temperatures, but it comes with a cost: a significant expenditure of evaporative water. This cost is important because it is found to increase with body size in Passeriformes due to the forced evaporative heat loss, but it shows virtually no increase with body size in Non-Passeriformes. Thus, despite a high BMR significantly increasing ecological opportunities, this way of expanding the ecological niches is possible for the small size class only. These findings suggest that the high level of basal metabolic rate in Passeriformes in comparison to Non-Passeriformes determines the necessity for the former to utilize considerably larger amounts of water for evaporation to maintain the needed heat balance, especially at higher ambient temperatures and at larger body sizes.

Ecological and scaling analysis of the energy expenditure of rest, activity, flight, and evaporative water loss in Passeriformes and non-Passeriformes in relation to seasonal migrations and to the occupation of boreal stations in high and moderate latitudes.

A unified system of bioenergetic parameters that describe thermal regulation and energy metabolism in many passerine and non-passerine species has been developed. These parameters have been analyzed as functions of ambient temperature, and bioenergetic models for various species have been developed. The level of maximum food energy or maximal existence metabolism (MPE) is 1.3 times higher in passerines than in non-passerines, which is consistent with the ratio of their basal metabolic rates (BMR). The optimal ambient temperature for maximizing productive processes (e.g., reproduction, molting) is lower for passerines than for non passerines, which allows passerines to have higher production rates at moderate ambient temperatures. This difference in the optimal ambient temperature may explain the variation in bioenergetic parameters along latitudinal gradients, such as the well-known ecological rule of clutch size (or mass) increase in the more northerly passerine birds. The increased potential for productive energy output in the north may also allow birds to molt faster there. This phenomenon allows passerine birds to occupy a habitat that fluctuates widely in ambient temperature compared with non-passerine birds of similar size. Passerines have a more effective system for maintaining heat balance at both high and low temperatures. The high metabolism and small body sizes of passerines are consistent with omnivore development and with ecological plasticity. Among large passerines, the unfavorable ratio of MPE to BMR should decrease the energy that is available for productive processes. This consequence limits both the reproductive output and the development of long migration (particularly in Corvus corax). The hypothesis regarding BMR increase in passerines was suggested based on an aerodynamic analysis of the flight speed and the wing characteristics. This allometric analysis shows that the flight velocity is approximately 20% lower in Passeriformes than in non-Passeriformes, which is consistent with the inverted ratio of their BMR level. The regressions for the aerodynamic characteristics of wings show that passerines do not change the morphological characteristics of their wings to decrease velocity. Passerine birds prefer forest habitats. The size range of 5-150 g for birds in forest habitats is almost exclusively occupied by passerines because of their large energetic capability.

Mean mass-specific metabolic rates are strikingly similar across life's major domains: Evidence for life's metabolic optimum.

A fundamental but unanswered biological question asks how much energy, on average, Earth's different life forms spend per unit mass per unit time to remain alive. Here, using the largest database to date, for 3,006 species that includes most of the range of biological diversity on the planet-from bacteria to elephants, and algae to sapling trees-we show that metabolism displays a striking degree of homeostasis across all of life. We demonstrate that, despite the enormous biochemical, physiological, and ecological differences between the surveyed species that vary over 10(20)-fold in body mass, mean metabolic rates of major taxonomic groups displayed at physiological rest converge on a narrow range from 0.3 to 9 W kg(-1). This 30-fold variation among life's disparate forms represents a remarkably small range compared with the 4,000- to 65,000-fold difference between the mean metabolic rates of the smallest and largest organisms that would be observed if life as a whole conformed to universal quarter-power or third-power allometric scaling laws. The observed broad convergence on a narrow range of basal metabolic rates suggests that organismal designs that fit in this physiological window have been favored by natural selection across all of life's major kingdoms, and that this range might therefore be considered as optimal for living matter as a whole.