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Examines thermoregulation in the flowers of the sacred lotus, Nelumbo nucifera. Temperature measurements of the flowers and surrounding air; Rates of oxygen consumption.

Presents a study on the influence of ambient temperature on thermoregulation by the protogynous inflorescence of eastern skunk cabbage, Symplocarpus foetidus, of the Araceae arum lily family. Recording of oxygen consumption, carbon dioxide production and temperature of plants; Bimodial distribution of temperatures due to switching between warm and cool states.

Analyzes the spadix temperature, rate of oxygen consumption and pattern of respiration in whole inflorescences attached to thermogenic arum Philodendron selloum. Effects of ambient temperature; Temperature regulation in peak and plateau phases; Respiratory rates and energetics; Role of thermogenesis in pollination.

A prevailing assumption in the cardiovascular field is that the metabolic rate of the heart is a constant proportion of a mammal's whole-body aerobic metabolic rate. In this Commentary, we assemble previously published cardiovascular, metabolic and body mass data from matched terrestrial mammalian species, at rest and during heavy exercise, and reveal scaling relationships that challenge this assumption. Our analyses indicate that the fractional metabolic cost of systemic perfusion compared with whole-body metabolic rate increases significantly with body size among resting mammals, from ∼2.5% in a mouse to ∼10% in an elephant. We propose that two significant body size-dependent effects contribute to this conclusion; namely, that larger species generate higher mean systemic arterial blood pressure and that their myocardium operates with lower external mechanical efficiencies compared with those of smaller species. We discuss potential physiological and mechanical explanations, including the additional energy needed to support the arterial blood column above the heart in larger species, especially those with long necks, as well as the possible sources of greater internal energy losses from the heart of larger species. Thus, we present an updated view of how increasing blood pressure and decreasing efficiency of the myocardium result in an increasing fractional metabolic cost of perfusion as body size increases among resting mammals., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

The whole-body (tachymetabolic) endothermy seen in modern birds and mammals is long held to have evolved independently in each group, a reasonable assumption when it was believed that its earliest appearances in birds and mammals arose many millions of years apart. That assumption is consistent with current acceptance that the non-shivering thermogenesis (NST) component of regulatory body heat originates differently in each group: from skeletal muscle in birds and from brown adipose tissue (BAT) in mammals. However, BAT is absent in monotremes, marsupials, and many eutherians, all whole-body endotherms. Indeed, recent research implies that BAT-driven NST originated more recently and that the biochemical processes driving muscle NST in birds, many modern mammals and the ancestors of both may be similar, deriving from controlled 'slippage' of Ca 2+ from the sarcoplasmic reticulum Ca 2+ -ATPase (SERCA) in skeletal muscle, similar to a process seen in some fishes. This similarity prompted our realisation that the capacity for whole-body endothermy could even have pre-dated the divergence of Amniota into Synapsida and Sauropsida, leading us to hypothesise the homology of whole-body endothermy in birds and mammals, in contrast to the current assumption of their independent (convergent) evolution. To explore the extent of similarity between muscle NST in mammals and birds we undertook a detailed review of these processes and their control in each group. We found considerable but not complete similarity between them: in extant mammals the 'slippage' is controlled by the protein sarcolipin (SLN), in birds the SLN is slightly different structurally and its role in NST is not yet proved. However, considering the multi-millions of years since the separation of synapsids and diapsids, we consider that the similarity between NST production in birds and mammals is consistent with their whole-body endothermy being homologous. If so, we should expect to find evidence for it much earlier and more widespread among extinct amniotes than is currently recognised. Accordingly, we conducted an extensive survey of the palaeontological literature using established proxies. Fossil bone histology reveals evidence of sustained rapid growth rates indicating tachymetabolism. Large body size and erect stature indicate high systemic arterial blood pressures and four-chambered hearts, characteristic of tachymetabolism. Large nutrient foramina in long bones are indicative of high bone perfusion for rapid somatic growth and for repair of microfractures caused by intense locomotion. Obligate bipedality appeared early and only in whole-body endotherms. Isotopic profiles of fossil material indicate endothermic levels of body temperature. These proxies led us to compelling evidence for the widespread occurrence of whole-body endothermy among numerous extinct synapsids and sauropsids, and very early in each clade's family tree. These results are consistent with and support our hypothesis that tachymetabolic endothermy is plesiomorphic in Amniota. A hypothetical structure for the heart of the earliest endothermic amniotes is proposed. We conclude that there is strong evidence for whole-body endothermy being ancient and widespread among amniotes and that the similarity of biochemical processes driving muscle NST in extant birds and mammals strengthens the case for its plesiomorphy., (© 2021 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

This meta-study uses phylogenetic scaling models across more than 30 species, spanning five orders of magnitude in body mass, to show that cardiac capillary numerical density and mitochondrial volume density decrease with body mass raised to the -0.07 ± 0.03 and -0.04 ± 0.01 exponents, respectively. Thus, while an average 10 g mammal has a cardiac capillary density of approximately 4150 mm -2 and a mitochondrial density of 33%, a 1 t mammal has considerably lower corresponding values of 1850 mm -2 and 21%. These similar scaling trajectories suggest quantitative matching for the primary oxygen supply and oxygen consuming structures of the heart, supporting economic design at the cellular level of the oxygen cascade in this aerobic organ. These scaling trajectories are nonetheless somewhat shallower than the exponent of -0.11 calculated for the maximum external mechanical power of the cardiac tissue, under conditions of heavy exercise, when oxygen flow between capillaries and mitochondria is probably fully exploited. This mismatch, if substantiated, implies a declining external mechanical efficiency of the heart with increasing body mass, whereby larger individuals put more energy in but get less energy out, a scenario with implications for cardiovascular design, aerobic capacity and limits of body size.

If arteries penetrate bones through foramina, regional artery blood flow rates can be estimated from the foramen sizes. Femoral bone blood flow rates estimated from nutrient foramen sizes were previously not absolute, but only a relative blood flow index (Q i ), because the size relationship between the foramen and the occupying artery was unknown. The current study used vascular contrast and micro-computerized tomographic scanning to investigate femoral nutrient foramen and nutrient artery sizes in three groups of sub-adult chickens (non-laying hens, laying hens, and roosters) of similar ages. The results indicate that the cross-sectional area of the nutrient artery lumen occupies approximately 20.2 ± 4.1% of the foramen for femora with only one foramen. Artery lumen size is significantly correlated with foramen size. Vascular contrast imaging is capable of estimating blood flow rates through nutrient arteries, as blood flow rates estimated from artery lumen casts are similar to blood flow rates measured by infusion of fluorescent-labeled microspheres. Laying hens tend to have higher nutrient artery perfusion rates than non-laying hens, probably due to extra oxygen and calcium requirements for eggshell production, although the calculated blood flow difference was not statistically significant. Histological embedding and sectioning along with vascular contrast imaging reveal variable nutrient foramen morphology and nutrient artery location among femora with more than one nutrient foramen., (© 2021 Anatomical Society.)

Respiratory gas exchange in avian embryos progresses through three stages inside the egg. During the first 3-5 days of incubation, the chicken embryo has no specialised respiratory organs and is not reliant on blood circulation. At this stage, it obtains oxygen mainly by diffusion through the eggshell, albumen, amniotic fluid and embryonic tissues. In the second stage, gas exchange relies on diffusion through the shell in the gas phase and convection by blood circulation through the chorioallantoic membrane and body. Day 19 starts the third stage, the transition from chorioallantoic to pulmonary gas exchange, which is complete when the chick hatches on day 20. Metabolism is thought to be aerobic throughout incubation, although the early embryo is covered by fluids (albumen and amniotic fluid) which would greatly resist oxygen diffusion. This study uses fibre-optic sensors to measure oxygen partial pressure (PO 2 ) near, and inside of, the embryo during days 3-5, and relates the data to total body lactate levels. The study shows that fluids surrounding the embryo greatly impede oxygen diffusion, with PO 2 becoming severely hypoxic near the embryo, occasionally almost anoxic inside it. Meanwhile, lactate rises to high levels, and the stored lactate can be later oxidised by the embryo when the chorioallantois takes over and metabolism becomes entirely aerobic., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The relationship between body mass and the respiratory microenvironment of burrowing animals is examined using artificial burrows containing surrogate animals that simulate O 2 consumption by removal of air and simultaneous replacement with N 2 . Allometric relationships between body mass and burrow radius, nest chamber radius, and O 2 consumption rate show that published mathematical predictions of diffusion-mediated gas exchange are adequate to describe the respiratory environments of animals in small blind-ending burrows through porous substrata. Diffusion is sufficient to ventilate burrows containing small mammals weighing less than 340 g, or subterranean nest chambers connected to the surface by one or more tunnels containing mammals weighing less than 30 kg. Outside of these limits, convection prevails and prevents the development of hypoxic conditions, particularly in burrows of mammals weighing more than 1300 g., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The metabolic rate of vertebrate bone tissue is related to bone growth, repair and homeostasis, which are all dependent on life stage. Bone metabolic rate is difficult to measure directly, but absolute blood flow rate () should reflect local tissue oxygen requirements. A recent 'foramen technique' has derived an index of blood flow rate () by measuring nutrient foramen sizes of long bones. is assumed to be proportional to ; however, the assumption has never been tested. This study used fluorescent microsphere infusion to measure femoral bone in anaesthetized non-laying hens, laying hens and roosters. Mean mass-specific cardiac output was 338±38 ml min-1 kg-1, and the two femora received 0.63±0.10% of this. Laying hens had higher wet bone mass-specific to femora (0.23±0.09 ml min-1 g-1) than the non-laying hens (0.12±0.06 ml min-1 g-1) and roosters (0.14±0.04 ml min-1 g-1), presumably associated with higher bone calcium mobilization during eggshell production. Estimated metabolic rate of femoral bone was 0.019 ml O2 min-1 g-1. Femoral increased significantly with body mass, but was not correlated with nutrient foramen radius (r), probably because of a narrow range in foramen radius. Over all 18 chickens, femoral shaft was 1.07±0.30 ml min-1 mm-1. Mean in chickens was significantly higher than predicted by an allometric relationship for adult cursorial bird species, possibly because the birds were still growing., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Many aquatic insects use bubbles on the body surface to store and supply O 2 for their dives. There are two types of bubbles: air stores, which store O 2 gained from air at the surface, and gas gills that allow passive extraction of O 2 from water. Many insects using air stores and gas gills return to the surface to replenish their bubbles and, therefore, their requirement for O 2 influences dive behaviour. In this study, we investigate gas exchange and dive behaviour in the diving beetle Platynectes decempunctatus that uses a sub-elytral air store and a small compressible gas gill. We measure the PO 2 within the air store during tethered dives, as well as the amount of O 2 exchanged during surfacing events. Buoyancy experiments monitor the volume of gas in the gas gill and how it changes during dives. We also directly link O 2 -consumption rate at three temperatures (10, 15 and 20 °C) with dive duration, surfacing frequency and movement activity. These data are incorporated in a gas exchange model, which shows that the small gas gill of P. decempunctatus contributes less than 10% of the total O 2 used during the dive, while up to 10% is supplied by cutaneous uptake., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

As an animal grows, the relative sizes of their organs may grow proportionately or disproportionately, depending on ontogenetic changes in function. If organ growth is proportional (isometric), then the exponent of the scaling equation is 1.0. Relative decreases or increases in size result in exponents less than 1 (hypoallometric) or greater than 1 (hyperallometric). Thus, the empirical exponent can indicate potential changes in function. The entire gastrointestinal tract (GIT) of the foregut-fermenting western grey kangaroo Macropus fuliginosus melanops exhibited biphasic allometry across five orders of magnitude body mass (M b ; 52.0 g-70.5 kg). Prior to weaning at around 12 kg M b , the entire empty GIT mass scaled with hyperallometry (M b 1.13 ), shifting to hypoallometry (M b 0.80 ) post-weaning. In addition, there were varying patterns of hyper-, hypo-, and isometric scaling for select GIT organs, with several displaying phase shifts associated with major life-history events, specifically around exit from the maternal pouch and around weaning. Mass of the kangaroo forestomach, the main fermentation site, scaled with hyperallometry (M b 1.16 ) before the stage of increased vegetation intake, and possibly after this stage (M b 1.12 ; P = 0.07), accompanied by a higher scaling factor (elevation of the curve) probably associated with more muscle for processing fibrous vegetation. The acid hindstomach mass showed hyperallometry (M b 1.15 ) before weaning, but hypoallometry (M b 0.58 ) post-weaning, presumably associated with decreasing intake of milk. Small intestine mass and length each scaled isometrically throughout ontogeny, with no discernible breakpoints at any life-history stage. The caecum and colon mass were steeply hyperallometric early in-pouch life (M b 1.59-1.66 ), when the young were ectothermic, hairless, and supported solely by milk. After around 295 g M b , caecum mass remained hyperallometric (M b 1.14 ), possibly supporting its early development as a nidus for microbial populations to provide for secondary fermentation in this organ after the young transition from milk to vegetation.

Blood flow rate ( Q ˙ ) in relation to arterial lumen radius (r i ) is commonly modelled according to theoretical equations and paradigms, including Murray's Law ( Q ˙ ∝ r i 3 ) and da Vinci's Rule ( Q ˙ ∝ r i 2 ). Wall shear stress (τ) is independent of r i with Murray's Law (τ ∝  r i 0 ) and decreases with da Vinci's Rule (τ ∝  r i - 1 ). These paradigms are tested empirically with a meta-analysis of the relationships between Q ˙ and r i in seven major arteries of the human cephalic circulation from 19 imaging studies in which both variables were presented. The analysis shows that Q ˙ ∝ r i 2.16 and τ ∝  r i - 1.02 , more consistent with da Vinci's Rule than Murray's Law. This meta-analysis provides standard values for Q ˙ , r i and τ in the human cephalic arteries that may be a useful baseline in future investigations. On average, the paired internal carotid arteries supply 75%, and the vertebral arteries supply 25%, of total brain blood flow. The internal carotid arteries contribute blood entirely to the anterior and middle cerebral arteries and also partly to the posterior cerebral arteries via the posterior communicating arteries of the circle of Willis. On average, the internal carotid arteries provide 88% of the blood flow to the cerebrum and the vertebral arteries only 12%., (© 2019 Anatomical Society.)

Some blood vessels enter bones through foramina, leaving the size of the foramen as a gauge for estimating the rate of blood flow and hence the metabolic rate of the supplied tissues. Foramen dimensions have been measured using varied methods in previous foramen studies, to relate regional blood flows with associated physiological processes. With the increasing interests in this 'foramen technique', standard methods with minimized measurement errors are therefore required. This study provides details of microphotographic and micro-computerized tomographic methods, and introduces a new alternative method, which uses impression material to measure foramen dimensions. The three methods are compared and the results indicate that all of them are capable of obtaining precise and accurate foramen dimension values, although they all have limitations. A microphotograph of the external opening is suggested to be the standard method because of its ease of use, but the alternative methods provide more detailed information on foramen shape. If the foramen is mainly occupied by one artery, blood flow rates can be calculated from foramen size and artery wall-lumen ratio, which is evaluated from the literature survey in this study. If veins or nerves also penetrate the foramen, a relative index of blood flow rate is nevertheless possible for comparative purposes., (© 2019 Anatomical Society.)

Brain metabolic rate (MR) is linked mainly to the cost of synaptic activity, so may be a better correlate of cognitive ability than brain size alone. Among primates, the sizes of arterial foramina in recent and fossil skulls can be used to evaluate brain blood flow rate, which is proportional to brain MR. We use this approach to calculate flow rate in the internal carotid arteries ( Q ˙ ICA ) , which supply most of the primate cerebrum. Q ˙ ICA is up to two times higher in recent gorillas, chimpanzees and orangutans compared with 3-million-year-old australopithecine human relatives, which had equal or larger brains. The scaling relationships between Q ˙ ICA and brain volume ( V br ) show exponents of 1.03 across 44 species of living haplorhine primates and 1.41 across 12 species of fossil hominins. Thus, the evolutionary trajectory for brain perfusion is much steeper among ancestral hominins than would be predicted from living primates. Between 4.4-million-year-old Ardipithecus and Homo sapiens , V br increased 4.7-fold, but Q ˙ ICA increased 9.3-fold, indicating an approximate doubling of metabolic intensity of brain tissue. By contrast, Q ˙ ICA is proportional to V br among haplorhine primates, suggesting a constant volume-specific brain MR.

Novel phenotypes are often linked to major ecological transitions during evolution. Here, we describe for the first time an unusual network of large blood vessels in the head of the sea snake Hydrophis cyanocinctus . MicroCT imaging and histology reveal an intricate modified cephalic vascular network (MCVN) that underlies a broad area of skin between the snout and the roof of the head. It is mostly composed of large veins and sinuses and converges posterodorsally into a large vein (sometimes paired) that penetrates the skull through the parietal bone. Endocranially, this blood vessel leads into the dorsal cerebral sinus, and from there, a pair of large veins depart ventrally to enter the brain. We compare the condition observed in H. cyanocinctus with that of other elapids and discuss the possible functions of this unusual vascular network. Sea snakes have low oxygen partial pressure in their arterial blood that facilitates cutaneous respiration, potentially limiting the availability of oxygen to the brain. We conclude that this novel vascular structure draining directly to the brain is a further elaboration of the sea snakes' cutaneous respiratory anatomy, the most likely function of which is to provide the brain with an additional supply of oxygen., Competing Interests: The authors have no competing interests., (© 2019 The Authors.)

Scaling of the heart across development can reveal the degree to which variation in cardiac morphology depends on body mass. In this study, we assessed the scaling of heart mass, left and right ventricular masses, and ventricular mass ratio, as a function of eviscerated body mass across fetal and postnatal development in Horro sheep Ovis aries (~50-fold body mass range; N = 21). Whole hearts were extracted from carcasses, cleaned, dissected into chambers and weighed. We found a biphasic relationship when heart mass was scaled against body mass, with a conspicuous 'breakpoint' around the time of birth, manifest not by a change in the scaling exponent (slope), but rather a jump in the elevation. Fetal heart mass (g) increased with eviscerated body mass (M b , kg) according to the power equation 4.90 M b 0.88 ± 0.26 (± 95% CI ) , whereas postnatal heart mass increased according to 10.0 M b 0.88 ± 0.10 . While the fetal and postnatal scaling exponents are identical (0.88) and reveal a clear dependence of heart mass on body mass, only the postnatal exponent is significantly less than 1.0, indicating the postnatal heart becomes a smaller component of body mass as the body grows, which is a pattern found frequently with postnatal cardiac development among mammals. The rapid doubling in heart mass around the time of birth is independent of any increase in body mass and is consistent with the normalization of wall stress in response to abrupt changes in volume loading and pressure loading at parturition. We discuss variation in scaling patterns of heart mass across development among mammals, and suggest that the variation results from a complex interplay between hard-wired genetics and epigenetic influences., (© 2019 Anatomical Society.)

Insects have a gas-filled respiratory system, which provides a challenge for those that have become aquatic secondarily. Diving beetles (Dytiscidae) use bubbles on the surface of their bodies to supply O 2 for their dives and passively gain O 2 from the water. However, these bubbles usually require replenishment at the water's surface. A highly diverse assemblage of subterranean dytiscids has evolved in isolated calcrete aquifers of Western Australia with limited/no access to an air-water interface, raising the question of how they are able to respire. We explored the hypothesis that they use cutaneous respiration by studying the mode of respiration in three subterranean dytiscid species from two isolated aquifers. The three beetle species consume O 2 directly from the water, but they lack structures on their bodies that could have respiratory function. They also have a lower metabolic rate than other insects. O 2 boundary layers surrounding the beetles are present, indicating that O 2 diffuses into the surface of their bodies via cutaneous respiration. Cuticle thickness measurements and other experimental results were incorporated into a mathematical model to understand whether cutaneous respiration limits beetle size. The model indicates that the cuticle contributes considerably to resistance in the O 2 cascade. As the beetles become larger, their metabolic scope narrows, potentially limiting their ability to allocate energy to mating, foraging and development at sizes above approximately 5 mg. However, the ability of these beetles to utilise cutaneous respiration has enabled the evolution of the largest assemblage of subterranean dytiscids in the world., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

This meta-study investigated the relationships between blood flow rate ( Q̇ ; cm 3  s -1 ), wall shear stress (τ w ; dyn cm -2 ) and lumen radius ( r i ; cm) in 20 named systemic arteries of nine species of mammals, ranging in mass from 23 g mice to 652 kg cows, at rest. In the dataset, derived from 50 studies, lumen radius varied between 3.7 µm in a cremaster artery of a rat and 11.2 mm in the aorta of a human. The 92 logged data points of [Formula: see text] and r i are described by a single second-order polynomial curve with the equation: [Formula: see text] The slope of the curve increased from approximately 2 in the largest arteries to approximately 3 in the smallest ones. Thus, da Vinci's rule ([Formula: see text]) applies to the main arteries and Murray's law ([Formula: see text]) applies to the microcirculation. A subset of the data, comprising only cephalic arteries in which [Formula: see text] is fairly constant, yielded the allometric power equation: [Formula: see text] These empirical equations allow calculation of resting perfusion rates from arterial lumen size alone, without reliance on theoretical models or assumptions on the scaling of wall shear stress in relation to body mass. As expected, [Formula: see text] of individual named arteries is strongly affected by body mass; however, [Formula: see text] of the common carotid artery from six species (mouse to horse) is also sensitive to differences in whole-body basal metabolic rate, independent of the effect of body mass., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Arum concinnatum Schott is a highly thermogenic species, with the temperature of the appendix exceeding ~10.9°C above the ambient temperature during thermogenesis, whereas the rates of respiration of the male florets in intact inflorescences peak at 0.92μmol s-1 g-1, which is the highest rate so far measured among the plants. Here, we attempt the ex situ exogenous induction of thermogenesis in whole inflorescences and in separate appendices of the spadix, and explore the thermogenic patterns under controlled laboratory conditions of light and temperature. Mature but unopened inflorescences and appendices showed thermogenic responses when treated with salicylic acid (SA), but not when treated with distilled water (control). With regard to light conditions, the responses revealed only one significant difference for inflorescences, which concerns the higher maximum temperature in the continuous light treatment compared with continuous dark. Along the ambient temperature gradient, at the lowest temperature edge individuals remained stable close to ambient temperature and to control. These findings suggest that, in general, ex situ exogenous induction of thermogenesis can be achieved in whole inflorescences and in separate appendices of spadix of A. concinnatum using SA. This study also indicates that SA acts independently of light conditions, while exogenous induction of thermogenesis takes place within an ambient temperature range.

Chemical immobilization is necessary for the physiological study of large wild animals. However, the immobilizing drugs can adversely affect the cardiovascular and respiratory systems, yielding data that do not accurately represent the normal, resting state. We hypothesize that these adverse effects can be ameliorated by reversing the immobilizing agent while holding the animal under general anaesthesia. We used habituated sheep Ovis aries (N = 5, 46.9 ± 5.3 kg body mass, mean ± SEM) and goats Capra hircus (N = 4, 27.7 ± 2.8 kg) as ungulate models for large wild animals, and measured their cardiorespiratory function under three conditions: (1) mild sedation (midazolam), as a proxy for the normal resting state, (2) immobilization (etorphine and azaperone), and (3) general anaesthesia (propofol) followed by etorphine antagonism (naltrexone). Cardiac output for both sheep and goats remained unchanged across the three conditions (overall means of 6.2 ± 0.9 and 3.3 ± 0.3 L min -1 , respectively). For both sheep and goats, systemic and pulmonary mean arterial pressures were significantly altered from initial midazolam levels when administered etorphine + azaperone, but those arterial pressures were restored upon transition to propofol anaesthesia and antagonism of the etorphine. Under etorphine + azaperone, minute ventilation decreased in the sheep, though this decrease was corrected under propofol, while the minute ventilation in the goats remained unchanged throughout. Under etorphine + azaperone, both sheep and goats displayed arterial blood hypoxia and hypercapnia (relative to midazolam levels), which failed to completely recover under propofol, indicating that more time might be needed for the blood gases to be adequately restored. Nonetheless, many of the confounding cardiorespiratory effects of etorphine were ameliorated when it was antagonized with naltrexone while the animal was held under propofol, indicating that this procedure can largely restore the cardiovascular and respiratory systems closer to a normal, resting state.

The hearts of smaller mammals tend to operate at higher mass-specific mechanical work rates than those of larger mammals. The ultrastructural characteristics of the heart that allow for such variation in work rate are still largely unknown. We have used perfusion-fixation, transmission electron microscopy and stereology to assess the morphology and anatomical aerobic power density of the heart as a function of body mass across six species of wild African antelope differing by approximately 20-fold in body mass. The survival of wild antelope, as prey animals, depends on competent cardiovascular performance. We found that relative heart mass (g kg -1 body mass) decreases with body mass according to a power equation with an exponent of -0.12±0.07 (±95% confidence interval). Likewise, capillary length density (km cm -3 of cardiomyocyte), mitochondrial volume density (fraction of cardiomyocyte) and mitochondrial inner membrane surface density (m 2  cm -3 of mitochondria) also decrease with body mass with exponents of -0.17±0.16, -0.06±0.05 and -0.07±0.05, respectively, trends likely to be associated with the greater mass-specific mechanical work rate of the heart in smaller antelope. Finally, we found proportionality between quantitative characteristics of a structure responsible for the delivery of oxygen (total capillary length) and those of a structure that ultimately uses that oxygen (total mitochondrial inner membrane surface area), which provides support for the economic principle of symmorphosis at the cellular level of the oxygen cascade in an aerobic organ., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

The aquatic bug Aphelocheirus aestivalis (Fabricius 1794) utilises a plastron, a thin bubble layer on the surface of its body to extract O 2 from the water. Millions of tiny hairs keep the bubble from collapsing, enabling the bug to remain submerged indefinitely. The development of fibre optic O 2 -probes has allowed measurements of O 2 pressure (PO 2 ) surrounding the plastron, and within the plastron although only for short periods. Here we developed methods to continuously measure plastron PO 2 , and investigate how it is affected by temperature (15, 20, 25°C), activity, and water circulation. We also made measurements of water PO 2 , temperature and velocity in the field and swimming velocity at the treatment temperatures. Results show that plastron PO 2 is inversely related to temperature, associated with differences in metabolic demand, and that small bouts of activity or changes in water convection result in rapid changes in plastron PO 2 . A model was developed to calculate the conditions under which Aphelocheirus would exist without becoming O 2 -limited in relation to water temperature, PO 2 and boundary layer thickness. This suggests that Aphelocheirus at one of two field sites may have a reduced metabolic scope even in well convected water in association with low PO 2 and moderate temperature, and that in well convected, air-saturated water, bugs may have a reduced metabolic scope where water temperatures are between 20 and 25°C. If exposed to 5kPa PO 2 , Aphelocheirus cannot sustain resting metabolic rate even in well-convected water and would die at temperatures above approximately 25°C., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

The nutrient artery passes through the nutrient foramen on the shaft of the femur and supplies more than half of the total blood flow to the bone. Assuming that the size of the nutrient foramen correlates with the size of the nutrient artery, an index of blood flow rate ( Q i ) can be calculated from nutrient foramen dimensions. Interspecific Q i is proportional to locomotor activity levels in adult mammals, birds and reptiles. However, no studies have yet estimated intraspecific Q i to test for the effects of growth and locomotor development on bone blood flow requirements. In this study, we used micro-CT and medical CT scanning to measure femoral dimensions and foramen radius to calculate femoral Q i during the in-pouch and post-pouch life stages of western grey kangaroos ( Macropus fuliginosus ) weighing 5.7 g to 70.5 kg and representing a 12,350-fold range in body mass. A biphasic scaling relationship between Q i and body mass was observed (breakpoint at ca. 1-5 kg body mass right before permanent pouch exit), with a steep exponent of 0.96±0.09 (95% CI) during the in-pouch life stage and a statistically independent exponent of -0.59±0.90 during the post-pouch life stage. In-pouch joeys showed Q i values that were 50-100 times higher than those of adult diprotodont marsupials of the same body mass, but gradually converged with them as post-pouch adults. Bone modelling during growth appears to be the main determinant of femoral bone blood flow during in-pouch development, whereas bone remodelling for micro-fracture repair due to locomotion gradually becomes the main determinant when kangaroos leave the pouch and become more active., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

The respiration rate of the thermogenic inflorescences of Japanese skunk cabbage Symplocarpus renifolius can reach 300 nmol s -1 g -1 , which is sufficient to raise spadix temperature (T s ) up to 15 °C above ambient air temperature (T a ). Respiration rate is inversely related to T a , such that the T s achieves a degree of independence from T a , an effect known as temperature regulation. Here, we measure oxygen consumption rate (Ṁo 2 ) in air (21% O 2 in mainly N 2 ) and in heliox (21% O 2 in He) to investigate the diffusive conductance of the network of gas-filled spaces and the thermoregulatory response. When T s was clamped at 15 °C, the temperature that produces maximal Ṁo 2 in this species, exposure to high diffusivity heliox increased mean Ṁo 2 significantly from 137 ± 17 to 202 ± 43 nmol s -1 g -1 FW, indicating that respiration in air is normally limited by diffusion in the gas phase and some mitochondria are unsaturated. When T a was clamped at 15 °C and T s was allowed to vary, exposure to heliox reduced T s 1 °C and increased Ṁo 2 significantly from 116 ± 10 to 137 ± 19 nmol s -1 g -1 , indicating that enhanced heat loss by conduction and convection can elicit the thermoregulatory response., (© 2017 John Wiley & Sons Ltd.)

Bipedal hopping is used by macropods, including rat-kangaroos, wallabies and kangaroos (superfamily Macropodoidea). Interspecific scaling of the ankle extensor muscle-tendon units in the lower hindlimbs of these hopping bipeds shows that peak tendon stress increases disproportionately with body size. Consequently, large kangaroos store and recover more strain energy in their tendons, making hopping more efficient, but their tendons are at greater risk of rupture. This is the first intraspecific scaling analysis on the functional morphology of the ankle extensor muscle-tendon units (gastrocnemius, plantaris and flexor digitorum longus) in one of the largest extant species of hopping mammal, the western grey kangaroo Macropus fuliginosus (5.8-70.5 kg post-pouch body mass). The effective mechanical advantage of the ankle extensors does not vary with post-pouch body mass, scaling with an exponent not significantly different from 0.0. Therefore, larger kangaroos balance rotational moments around the ankle by generating muscle forces proportional to weight-related gravitational forces. Maximum force is dependent upon the physiological cross-sectional area of the muscle, which we found scales geometrically with a mean exponent of only 0.67, rather than 1.0. Therefore, larger kangaroos are limited in their capacity to oppose large external forces around the ankle, potentially compromising fast or accelerative hopping. The strain energy return capacity of the ankle extensor tendons increases with a mean exponent of ~1.0, which is much shallower than the exponent derived from interspecific analyses of hopping mammals (~1.4-1.9). Tendon safety factor (ratio of rupture stress to estimated peak hopping stress) is lowest in the gastrocnemius (< 2), and it decreases with body mass with an exponent of -0.15, extrapolating to a predicted rupture at 160 kg. Extinct giant kangaroos weighing 250 kg could therefore not have engaged in fast hopping using 'scaled-up' lower hindlimb morphology of extant western grey kangaroos., (© 2017 Anatomical Society.)

Flying insects have the highest mass-specific metabolic rate of all animals. Oxygen is supplied to the flight muscles by a combination of diffusion and convection along the internal air-filled tubes of the tracheal system. This study measured maximum flight metabolic rate (FMR) during tethered flight in the migratory locust Locusta migratoria under varying oxygen partial pressure ( P O 2 ) in background gas mixtures of nitrogen (N 2 ), sulfur hexafluoride (SF 6 ) and helium (He), to vary O 2 diffusivity and gas mixture density independently. With N 2 as the sole background gas (normodiffusive-normodense), mass-independent FMR averaged 132±19 mW g -0.75 at normoxia ( P O 2 =21 kPa), and was not limited by tracheal system conductance, because FMR did not increase in hyperoxia. However, FMR declined immediately with hypoxia, oxy-conforming nearly completely. Thus, the locust respiratory system is matched to maximum functional requirements, with little reserve capacity. With SF 6 as the sole background gas (hypodiffusive-hyperdense), the shape of the relationship between FMR and P O 2  was similar to that in N 2 , except that FMR was generally lower (e.g. 24% lower at normoxia). This appeared to be due to increased density of the gas mixture rather than decreased O 2 diffusivity, because hyperoxia did not reverse it. Normoxic FMR was not significantly different in He-SF 6 (hyperdiffusive-normodense) compared with the N 2 background gas, and likewise there was no significant difference between FMR in SF 6 -He (normodiffusive-hyperdense) compared with the SF 6 background gas. The results indicate that convection, not diffusion, is the main mechanism of O 2 delivery to the flight muscle of the locust when demand is high., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

Due in part to their large size, aggressive temperament, and difficulty in handling, there are few physiological studies of adult crocodilians in the literature. As a result, studies comparing individuals across an ontogenetic series and comparisons among species are also lacking. We addressed this gap in knowledge by measuring standard metabolic rates (SMR) of three species of crocodilians (Crocodylus porosus, C. johnsoni, and Alligator mississippiensis), and included individuals that ranged from 0.22 to 114 kg. Allometric scaling of SMR with body mass was similar among the species, but C. porosus had significantly higher SMR than did C. johnsoni or A. mississippiensis. Differences in SMR among species are potentially related to behavioural differences in levels of aggression; C. porosus are the most aggressive of the crocodilians measured, and have rates of standard metabolism that are approximately 36% higher at the grand mean body size than those measured for C. johnsoni or A. mississippiensis, which are among the least aggressive crocodilians., Competing Interests: The authors have declared that no competing interests exist.

Cardiovascular function in dinosaurs can be inferred from fossil evidence with knowledge of how metabolic rate, blood flow rate, blood pressure, and heart size are related to body size in living animals. Skeletal stature and nutrient foramen size in fossil femora provide direct evidence of a high arterial blood pressure, a large four-chambered heart, a high aerobic metabolic rate, and intense locomotion. But was the heart of a huge, long-necked sauropod dinosaur able to pump blood up 9 m to its head?, (©2016 Int. Union Physiol. Sci./Am. Physiol. Soc.)

This study presents a structure-function analysis of the mammalian left ventricle and examines the performance of the cardiac capillary network, mitochondria, and myofibrils at rest and during simulated heavy exercise. Left ventricular external mechanical work rate was calculated from cardiac output and systemic mean arterial blood pressure in resting sheep (Ovis aries; n = 4) and goats (Capra hircus; n = 4) under mild sedation, followed by perfusion-fixation of the left ventricle and quantification of the cardiac capillary-tissue geometry and cardiomyocyte ultrastructure. The investigation was then extended to heavy exercise by increasing cardiac work according to published hemodynamics of sheep and goats performing sustained treadmill exercise. Left ventricular work rate averaged 0.017 W/cm 3 of tissue at rest and was estimated to increase to ∼0.060 W/cm 3 during heavy exercise. According to an oxygen transport model we applied to the left ventricular tissue, we predicted that oxygen consumption increases from 195 nmol O 2 ·s -1 ·cm -3 of tissue at rest to ∼600 nmol O 2 ·s -1 ·cm -3 during heavy exercise, which is within 90% of the oxygen demand rate and consistent with work remaining predominantly aerobic. Mitochondria represent 21-22% of cardiomyocyte volume and consume oxygen at a rate of 1,150 nmol O 2 ·s -1 ·cm -3 of mitochondria at rest and ∼3,600 nmol O 2 ·s -1 ·cm -3 during heavy exercise, which is within 80% of maximum in vitro rates and consistent with mitochondria operating near their functional limits. Myofibrils represent 65-66% of cardiomyocyte volume, and according to a Laplacian model of the left ventricular chamber, generate peak fiber tensions in the range of 50 to 70 kPa at rest and during heavy exercise, which is less than maximum tension of isolated cardiac tissue (120-140 kPa) and is explained by an apparent reserve capacity for tension development built into the left ventricle., (Copyright © 2016 the American Physiological Society.)

The evolution of human cognition has been inferred from anthropological discoveries and estimates of brain size from fossil skulls. A more direct measure of cognition would be cerebral metabolic rate, which is proportional to cerebral blood flow rate (perfusion). The hominin cerebrum is supplied almost exclusively by the internal carotid arteries. The sizes of the foramina that transmitted these vessels in life can be measured in hominin fossil skulls and used to calculate cerebral perfusion rate. Perfusion in 11 species of hominin ancestors, from Australopithecus to archaic Homo sapiens , increases disproportionately when scaled against brain volume (the allometric exponent is 1.41). The high exponent indicates an increase in the metabolic intensity of cerebral tissue in later Homo species, rather than remaining constant (1.0) as expected by a linear increase in neuron number, or decreasing according to Kleiber's Law (0.75). During 3 Myr of hominin evolution, cerebral tissue perfusion increased 1.7-fold, which, when multiplied by a 3.5-fold increase in brain size, indicates a 6.0-fold increase in total cerebral blood flow rate. This is probably associated with increased interneuron connectivity, synaptic activity and cognitive function, which all ultimately depend on cerebral metabolic rate.

Thermoregulation (homeothermy) in animals involves a complex mechanism involving thermal receptors throughout the body and integration in the hypothalamus that controls shivering and non-shivering thermogenesis. The flowers of some ancient families of seed plants show a similar degree of physiological thermoregulation, but by a different mechanism. Here, we show that respiratory control in homeothermic spadices of skunk cabbage (Symplocarpus renifolius) is achieved by rate-determining biochemical reactions in which the overall thermodynamic activation energy exhibits a negative value. Moreover, NADPH production, catalyzed by mitochondrial isocitrate dehydrogenase in a chemically endothermic reaction, plays a role in the pre-equilibrium reaction. We propose that a law of chemical equilibrium known as Le Châtelier's principle governs the homeothermic control in skunk cabbage.

In a recent technical comment regarding our analysis of the scaling of blood pressure with body mass in mammals (White and Seymour 2014), Packard (2015) argues that the trends in our graphs do not accurately reflect the relationship between the original variables, and that neither the graphics nor the accompanying statistical analyses provide strong support for the conclusions from the study, namely that larger mammals have higher arterial blood pressures. Here we take the opportunity to respond to these criticisms., (© 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.)

Many aquatic insects utilise air bubbles on the surface of their bodies to supply O2 while they dive. The bubbles can simply store O2, as in the case of an 'air store', or they can act as a physical 'gas gill', extracting O2 from the water. Backswimmers of the genus Anisops augment their air store with O2 from haemoglobin cells located in the abdomen. The O2 release from the haemoglobin helps stabilise bubble volume, enabling backswimmers to remain near neutrally buoyant for a period of the dive. It is generally assumed that the backswimmer air store does not act as a gas gill and that gas exchange with the water is negligible. This study combines measurements of dive characteristics under different exotic gases (N2, He, SF6, CO) with mathematical modelling, to show that the air store of the backswimmer Anisops deanei does exchange gases with the water. Our results indicate that approximately 20% of O2 consumed during a dive is obtained directly from the water. Oxygen from the water complements that released from the haemoglobin, extending the period of near-neutral buoyancy and increasing dive duration., (© 2015. Published by The Company of Biologists Ltd.)

The river bug Aphelocheirus aestivalis is a 40 mg aquatic insect that, as an adult, relies totally on an incompressible physical gill to exchange respiratory gases with the water. The gill (called a 'plastron') consists of a stationary layer of air held in place on the body surface by millions of tiny hairs that support a permanent air-water interface, so that the insect never has to renew the gas at the water's surface. The volume of air in the plastron is extremely small (0.14 mm(3)), under slightly negative pressure and connected to the gas-filled tracheal system through spiracles on the cuticle. Here, we measure PO2 of the water and within the plastron gas with O2-sensing fibre optics to understand the effectiveness and limitations of the gas exchanger. The difference in PO2 is highest in stagnant water and decreases with increasing convection over the surface. Respiration of bugs in water-filled vials varies between 33 and 296 pmol O2 s(-1), depending on swimming activity. The effective thickness of the boundary layer around the plastron was calculated from respiration rate, PO2 difference and plastron surface area, according to the Fick diffusion equation and verified by direct measurements with the fibre-optic probes. In stagnant water, the boundary layer is approximately 500 μm thick, which nevertheless can satisfy the demands of resting bugs, even if the PO2 of the free water decreases to half that of air saturation. Active bugs require thinner boundary layers (∼ 100 μm), which are achieved by living in moving water or by swimming., (© 2015. Published by The Company of Biologists Ltd.)

The evolution of primates involved increasing body size, brain size and presumably cognitive ability. Cognition is related to neural activity, metabolic rate and rate of blood flow to the cerebral cortex. These parameters are difficult to quantify in living animals. This study shows that it is possible to determine the rate of cortical brain perfusion from the size of the internal carotid artery foramina in skulls of certain mammals, including haplorrhine primates and diprotodont marsupials. We quantify combined blood flow rate in both internal carotid arteries as a proxy of brain metabolism in 34 species of haplorrhine primates (0.116-145 kg body mass) and compare it to the same analysis for 19 species of diprotodont marsupials (0.014-46 kg). Brain volume is related to body mass by essentially the same exponent of 0.70 in both groups. Flow rate increases with haplorrhine brain volume to the 0.95 power, which is significantly higher than the exponent (0.75) expected for most organs according to 'Kleiber's Law'. By comparison, the exponent is 0.73 in marsupials. Thus, the brain perfusion rate increases with body size and brain size much faster in primates than in marsupials. The trajectory of cerebral perfusion in primates is set by the phylogenetically older groups (New and Old World monkeys, lesser apes) and the phylogenetically younger groups (great apes, including humans) fall near the line, with the highest perfusion. This may be associated with disproportionate increases in cortical surface area and mental capacity in the highly social, larger primates., (© 2015. Published by The Company of Biologists Ltd.)

The heart and left ventricle of the marsupial western grey kangaroo Macropus fuliginosus exhibit biphasic allometric growth, whereby a negative shift in the trajectory of cardiac growth occurs at pouch exit. In this study, we used transmission electron microscopy to examine the scaling of left ventricle cardiomyocyte ultrastructure across development in the western grey kangaroo over a 190-fold body mass range (0.355-67.5 kg). The volume-density (%) of myofibrils, mitochondria, sarcoplasmic reticuli and T-tubules increase significantly during in-pouch growth, such that the absolute volume (ml) of these organelles scales with body mass (Mb; kg) with steep hyperallometry: 1.41Mb (1.38), 0.64Mb (1.29), 0.066Mb (1.45) and 0.035Mb (1.87), respectively. Maturation of the left ventricle ultrastructure coincides with pouch vacation, as organelle volume-densities scale independent of body mass across post-pouch development, such that absolute organelle volumes scale in parallel and with relatively shallow hypoallometry: 4.65Mb (0.79), 1.75Mb (0.77), 0.21Mb (0.79) and 0.35Mb (0.79), respectively. The steep hyperallometry of organelle volumes and volume-densities across in-pouch growth is consistent with the improved contractile performance of isolated cardiac muscle during fetal development in placental mammals, and is probably critical in augmenting cardiac output to levels necessary for endothermy and independent locomotion in the young kangaroo as it prepares for pouch exit. The shallow hypoallometry of organelle volumes during post-pouch growth suggests a decrease in relative cardiac requirements as body mass increases in free-roaming kangaroos, which is possibly because the energy required for hopping is independent of speed, and the capacity for energy storage during hopping could increase as the kangaroo grows., (© 2015. Published by The Company of Biologists Ltd.)