Measurement of microclimates in a warming world: problems and solutions.

As the world warms, it will be tempting to relate the biological responses of terrestrial animals to air temperature. But air temperature typically plays a lesser role in the heat exchange of those animals than does radiant heat. Under radiant load, animals can gain heat even when body surface temperature exceeds air temperature. However, animals can buffer the impacts of radiant heat exposure: burrows and other refuges may block solar radiant heat fully, but trees and agricultural shelters provide only partial relief. For animals that can do so effectively, evaporative cooling will be used to dissipate body heat. Evaporative cooling is dependent directly on the water vapour pressure difference between the body surface and immediate surroundings, but only indirectly on relative humidity. High relative humidity at high air temperature implies a high water vapour pressure, but evaporation into air with 100% relative humidity is not impossible. Evaporation is enhanced by wind, but the wind speed reported by meteorological services is not that experienced by animals; instead, the wind, air temperature, humidity and radiation experienced is that of the animal's microclimate. In this Commentary, we discuss how microclimate should be quantified to ensure accurate assessment of an animal's thermal environment. We propose that the microclimate metric of dry heat load to which the biological responses of animals should be related is black-globe temperature measured on or near the animal, and not air temperature. Finally, when analysing those responses, the metric of humidity should be water vapour pressure, not relative humidity.

Centralized red muscle in Odontaspis ferox and the prevalence of regional endothermy in sharks.

The order Lamniformes contains charismatic species such as the white shark Carcharodon carcharias and extinct megatooth shark Otodus megalodon, and is of particular interest given their influence on marine ecosystems, and because some members exhibit regional endothermy. However, there remains significant debate surrounding the prevalence and evolutionary origin of regional endothermy in the order, and therefore the development of phenomena such as gigantism and filter-feeding in sharks generally. Here we show a basal lamniform shark, the smalltooth sand tiger shark Odontaspis ferox, has centralized skeletal red muscle and a thick compact-walled ventricle; anatomical features generally consistent with regionally endothermy. This result, together with the recent discovery of probable red muscle endothermy in filter feeding basking sharks Cetorhinus maximus, suggests that this thermophysiology is more prevalent in the Lamniformes than previously thought, which in turn has implications for understanding the evolution of regional endothermy, gigantism, and extinction risk of warm-bodied shark species both past and present.

Relationship between capillaries, mitochondria and maximum power of the heart: a meta-study from shrew to elephant.

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.

Scaling at different ontogenetic stages: Gastrointestinal tract contents of a marsupial foregut fermenter, the western grey kangaroo Macropus fuliginosus melanops.

Prominent ontogenetic changes of the gastrointestinal tract (GIT) should occur in mammals whose neonatal diet of milk differs from that of adults, and especially in herbivores (as vegetation is particularly distinct from milk), and even more so in foregut fermenters, whose forestomach only becomes functionally relevant with vegetation intake. Due to the protracted lactation in marsupials, ontogenetic differences can be particularly well investigated in this group. Here, we report body mass (BM) scaling relationships of wet GIT content mass in 28 in-pouch young (50 g to 3 kg) and 15 adult (16-70 kg) western grey kangaroos Macropus fuliginosus melanops. Apart from the small intestinal contents, in-pouch young and adults did not differ in the scaling exponents ('slope' in log-log plots) but did differ in the scaling factor ('intercept'), with an implied substantial increase in wet GIT content mass during the out-of-pouch juvenile period. In contrast to forestomach contents, caecum contents were elevated in juveniles still in the pouch, suggestive of fermentative digestion of milk and intestinal secretion residues, particularly in the caecum. The substantial increase in GIT contents (from less than 1 to 10-20% of BM) was associated mainly with the increase in forestomach contents (from 25 to 80% of total GIT contents) and a concomitant decrease in small intestine contents (from 50 to 8%), emphasizing the shifting relevance of auto-enzymatic and allo-enzymatic (microbial) digestion. There was a concomitant increase in the contents-to-tissue ratio of the fermentation chambers (forestomach and caecum), but this ratio generally did not change for the small intestine. Our study not only documents significant ontogenetic changes in digestive morpho-physiology, but also exemplifies the usefulness of intraspecific allometric analyses for quantifying these changes.

Reliability of pulse oximetry at four different attachment sites in immobilized white rhinoceros (Ceratotherium simum).

OBJECTIVES: To determine the reliability of peripheral oxygen haemoglobin saturation (SpO2), measured by a Nonin PalmSAT 2500A pulse oximeter with 2000T transflectance probes at four attachment sites (third eyelid, cheek, rectum and tail), by comparing these measurements to arterial oxygen haemoglobin saturation (SaO2), measured by an AVOXimeter 4000 co-oximeter reference method in immobilized white rhinoceros (Ceratotherium simum). STUDY DESIGN: Randomized crossover study. ANIMALS: A convenience sample of eight wild-caught male white rhinoceros. METHODS: White rhinoceros were immobilized with etorphine (0.0026 ± 0.0002 mg kg-1, mean ± standard deviation) intramuscularly, after which the pinna was aseptically prepared for arterial blood sample collection, and four pulse oximeters with transflectance probes were fixed securely to their attachment sites (third eyelid, cheek, rectum and tail). At 30 minutes following recumbency resulting from etorphine administration, the animals were given either butorphanol (0.026 ± 0.0001 mg kg-1) or an equivalent volume of saline intravenously. At 60 minutes following recumbency, insufflated oxygen (15 L minute-1 flow rate) was provided intranasally. In total, the SpO2 paired measurements from the third eyelid (n = 80), cheek (n = 67), rectum (n = 59) and tail (n = 76) were compared with near-simultaneous SaO2 measurements using Bland-Altman to assess bias (accuracy), precision, and the area root mean squares (ARMS) method. RESULTS: Compared with SaO2, SpO2 measurements from the third eyelid were reliable (i.e., accurate and precise) above an SaO2 range of 70% (bias = 1, precision = 3, ARMS = 3). However, SpO2 measurements from the cheek, rectum and tail were unreliable (i.e., inaccurate or imprecise). CONCLUSIONS AND CLINICAL RELEVANCE: A Nonin PalmSAT pulse oximeter with a transflectance probe inserted into the space between the third eyelid and the sclera provided reliable SpO2 measurements when SaO2 was > 70%, in immobilized white rhinoceros.

How dryland mammals will respond to climate change: the effects of body size, heat load and a lack of food and water.

Mammals in drylands are facing not only increasing heat loads but also reduced water and food availability as a result of climate change. Insufficient water results in suppression of evaporative cooling and therefore increases in body core temperature on hot days, while lack of food reduces the capacity to maintain body core temperature on cold nights. Both food and water shortage will narrow the prescriptive zone, the ambient temperature range over which body core temperature is held relatively constant, which will lead to increased risk of physiological malfunction and death. Behavioural modifications, such as shifting activity between night and day or seeking thermally buffered microclimates, may allow individuals to remain within the prescriptive zone, but can incur costs, such as reduced foraging or increased competition or predation, with consequences for fitness. Body size will play a major role in predicting response patterns, but identifying all the factors that will contribute to how well dryland mammals facing water and food shortage will cope with increasing heat loads requires a better understanding of the sensitivities and responses of mammals exposed to the direct and indirect effects of climate change.

Blood flow rate and wall shear stress in seven major cephalic arteries of humans.

Blood flow rate ( Q˙ ) in relation to arterial lumen radius (ri ) is commonly modelled according to theoretical equations and paradigms, including Murray's Law ( Q˙ ∝ ri3 ) and da Vinci's Rule ( Q˙ ∝ ri2 ). Wall shear stress (τ) is independent of ri with Murray's Law (τ âˆ  ri0 ) and decreases with da Vinci's Rule (τ âˆ  ri-1 ). These paradigms are tested empirically with a meta-analysis of the relationships between Q˙ and ri in seven major arteries of the human cephalic circulation from 19 imaging studies in which both variables were presented. The analysis shows that Q˙ ∝ ri2.16 and τ âˆ  ri-1.02 , more consistent with da Vinci's Rule than Murray's Law. This meta-analysis provides standard values for Q˙ , ri 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%.

Functional morphology of the ankle extensor muscle-tendon units in the springhare Pedetes capensis shows convergent evolution with macropods for bipedal hopping locomotion.

This study assesses the functional morphology of the ankle extensor muscle-tendon units of the springhare Pedetes capensis, an African bipedal hopping rodent, to test for convergent evolution with the Australian bipedal hopping macropods. We dissect and measure the gastrocnemius, soleus, plantaris, and flexor digitorum longus in 10 adult springhares and compare them against similar-sized macropods using phylogenetically informed scaling analyses. We show that springhares align reasonably well with macropod predictions, being statistically indistinguishable with respect to the ankle extensor mean weighted muscle moment arm (1.63 vs. 1.65 cm, respectively), total muscle mass (41.1 vs. 29.2 g), total muscle physiological cross-sectional area (22.9 vs. 19.3 cm2 ), mean peak tendon stress (26.2 vs. 35.2 MPa), mean tendon safety factor (4.7 vs. 3.6), and total tendon strain energy return capacity (1.81 vs. 1.82 J). However, total tendon cross-sectional area is significantly larger in springhares than predicted for a similar-sized macropod (0.26 vs. 0.17 cm2 , respectively), primarily due to a greater plantaris tendon thickness (0.084 vs. 0.048 cm2 ), and secondarily because the soleus muscle-tendon unit is present in springhares but is vestigial in macropods. The overall similarities between springhares and macropods indicate that evolution has favored comparable lower hindlimb body plans for bipedal hopping locomotion in the two groups of mammals that last shared a common ancestor ~160 million years ago. The springhare's relatively thick plantaris tendon may facilitate rapid transfer of force from muscle to skeleton, enabling fast and accelerative hopping, which could help to outpace and outmaneuver predators.

Cerebral blood flow rates in recent great apes are greater than in Australopithecus species that had equal or larger brains.

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 (Vbr) 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, Vbr 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 Vbr among haplorhine primates, suggesting a constant volume-specific brain MR.

Scaling of cardiac morphology is interrupted by birth in the developing sheep Ovis aries.

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 (Mb , kg) according to the power equation 4.90 Mb0.88 ± 0.26 (± 95%CI) , whereas postnatal heart mass increased according to 10.0 Mb0.88 ± 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.

Interspecific scaling of blood flow rates and arterial sizes in mammals.

This meta-study investigated the relationships between blood flow rate (Q̇; cm3 s-1), wall shear stress (τw; dyn cm-2) and lumen radius (ri; 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 ri 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.

Femoral bone perfusion through the nutrient foramen during growth and locomotor development of western grey kangaroos (Macropus fuliginosus).

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 (Qi) can be calculated from nutrient foramen dimensions. Interspecific Qi is proportional to locomotor activity levels in adult mammals, birds and reptiles. However, no studies have yet estimated intraspecific Qi 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 Qi 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 Qi 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 Qi 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.

Scaling of morphology and ultrastructure of hearts among wild African antelope.

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 (m2 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.

Revisiting concepts of thermal physiology: Predicting responses of mammals to climate change.

The accuracy of predictive models (also known as mechanistic or causal models) of animal responses to climate change depends on properly incorporating the principles of heat transfer and thermoregulation into those models. Regrettably, proper incorporation of these principles is not always evident. We have revisited the relevant principles of thermal physiology and analysed how they have been applied in predictive models of large mammals, which are particularly vulnerable, to climate change. We considered dry heat exchange, evaporative heat transfer, the thermoneutral zone and homeothermy, and we examined the roles of size and shape in the thermal physiology of large mammals. We report on the following misconceptions in influential predictive models: underestimation of the role of radiant heat transfer, misassignment of the role and misunderstanding of the sustainability of evaporative cooling, misinterpretation of the thermoneutral zone as a zone of thermal tolerance or as a zone of sustainable energetics, confusion of upper critical temperature and critical thermal maximum, overestimation of the metabolic energy cost of evaporative cooling, failure to appreciate that the current advantages of size and shape will become disadvantageous as climate change advances, misassumptions about skin temperature and, lastly, misconceptions about the relationship between body core temperature and its variability with body mass in large mammals. Not all misconceptions invalidate the models, but we believe that preventing inappropriate assumptions from propagating will improve model accuracy, especially as models progress beyond their current typically static format to include genetic and epigenetic adaptation that can result in phenotypic plasticity.

Scaling of the ankle extensor muscle-tendon units and the biomechanical implications for bipedal hopping locomotion in the post-pouch kangaroo Macropus fuliginosus.

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.

Flight metabolic rate of Locusta migratoria in relation to oxygen partial pressure in atmospheres of varying diffusivity and density.

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 (PO2 ) in background gas mixtures of nitrogen (N2), sulfur hexafluoride (SF6) and helium (He), to vary O2 diffusivity and gas mixture density independently. With N2 as the sole background gas (normodiffusive-normodense), mass-independent FMR averaged 132±19 mW g-0.75 at normoxia (PO2 =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 SF6 as the sole background gas (hypodiffusive-hyperdense), the shape of the relationship between FMR and PO2  was similar to that in N2, 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 O2 diffusivity, because hyperoxia did not reverse it. Normoxic FMR was not significantly different in He-SF6 (hyperdiffusive-normodense) compared with the N2 background gas, and likewise there was no significant difference between FMR in SF6-He (normodiffusive-hyperdense) compared with the SF6 background gas. The results indicate that convection, not diffusion, is the main mechanism of O2 delivery to the flight muscle of the locust when demand is high.

Gas exchange and dive characteristics of the free-swimming backswimmer Anisops deanei.

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.

Scaling of cerebral blood perfusion in primates and marsupials.

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.

Scaling of left ventricle cardiomyocyte ultrastructure across development in the kangaroo Macropus fuliginosus.

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.