Metabolic Rate Activity Research Articles

Obesity Influence on Insulin Activity and Resting Metabolic Rate in Type 2 Diabetes

AbstractBackground and Aims. Diabetes mellitus and obesity modify the resting metabolic rate (RMR) in opposite directions. This study aimed to evaluate the relationship between RMR, insulin activity and adipocytokines in type 2 diabetes patients with associated obesity. Material and Method. Anthropometric and biochemical measurements, bioimpedance body composition, C peptide, insulin, proinsulin, adiponectin, leptin, indirect calorimetry measured RMR were determined in 306 (53% male, 54.9% obese) diabetic patients. Results. RMR, C peptide, insulin, proinsulin, Homeostasis Model Assessment (HOMA) IR, HOMA %B, leptin increased and adiponectin decreased with body mass index (BMI). BMI had a statistical significant higher effect size on fat mass (FM), leptin (0.652-0.339 eta squared), moderate positive effect size on fat-free mass (FFM), insulin, C peptide, RMR, HOMA %B (0.228 - 0.155 eta squared) and lower effect size on HOMA-IR, adiponectin, proinsulin-to-adiponectin ratio, proinsulin (0.098-0.062 eta squared). RMR had positive correlations with FFM (r=0.633), BMI (r=0.300), proinsulin-to-adiponectin ratio (r=0.218), HOMA1-IR (r=0.176), proinsulin (r=0.151), insulin (r=0.144), fasting plasma glucose (r=0.132), fat mass (r= 0.131), triglycerides (r = 0.119), proinsulin-to-insulin ratio (r= 0.117) and negative with age (r=-0.368), HDL cholesterol (r=-0.284), adiponectin(r=-0.282) and leptin (r=-0.178). Conclusions RMR was correlated with HOMA IR and insulin resistance biomarkers, but not with HOMA %B and C peptide.

Swimming speeds and metabolic rates of semi-captive juvenile lemon sharks (Negaprion brevirostris, Poey) estimated with acceleration biologgers

There is much interest in being able to quantify the swimming speeds and metabolic rates of wild aquatic animals such as sharks to develop bioenergetics models and evaluate the metabolic consequences of different stressors. This study sought to calibrate tri-axial acceleration biologgers (accelerometers) such that it would be possible to estimate swimming speeds and metabolic rates of semi-captive sharks in an enclosed natural mesocosm. Juvenile lemon sharks (Negaprion brevirostris, Poey; 60–75cm total length, 1–2kg) were equipped with accelerometers and swum at stepwise velocity increments in a Blazka-style swim tunnel respirometer using a critical swimming speed protocol. Metabolic rates and acceleration-derived metrics (overall dynamic body acceleration and tailbeat frequency) were measured concurrently during forced swimming, and accelerometer-equipped sharks were released to an enclosed mesocosm habitat to estimate average daily metabolic rate (ADMR) and swimming velocities across diel and tidal cycles. Acceleration-derived tailbeat frequency was a stronger predictor of metabolic rate than overall dynamic body action, and predicted an active metabolic rate of 249.7±1.9mg O2 kg−1h−1 and an ADMR of 88.7±0.7kJkg−1d−1 at 30°C. Following exhaustive exercise a maximum metabolic rate of 398.0±19.6mg O2 kg−1h−1 was achieved and over the subsequent 55-minute recovery period excess post-exercise oxygen consumption was 31.2mg O2 kg−1. The critical swimming speed of the sharks was 0.71±0.03 body lengths per second (BL s−1), and swimming speed in the mesocosm was 0.19±0.01 BL s−1. Locomotor activity levels of semi-captive sharks in the mesocosm were influenced by tide state and diel period, with sharks having highest swimming velocities during diurnal periods and flooding tides. Overall, accelerometry is a suitable means for estimating swimming speed and metabolic rate in this species, and additional research to address anaerobic energy expenditure of wild sharks is warranted.

Swimming metabolic rates vary by sex and development stage, but not by species, in three species of Australian otariid seals.

Physiology may limit the ability for marine mammals to adapt to changing environments. Depth and duration of foraging dives are a function of total available oxygen stores, which theoretically increase as animals grow, and metabolic costs. To evaluate how physiology may influence the travelling costs for seals to foraging patches in the wild, we measured metabolic rates of a cross-section of New Zealand fur seals, Australian fur seals and Australian sea lions representing different foraging strategies, development stages, sexes and sizes. We report values for standard metabolic rate, active metabolic rate (obtained from submerged swimming), along with estimates of cost of transport (COT), measured via respirometry. We found a decline in mass-specific metabolic rate with increased duration of submerged swimming. For most seals mass-specific metabolic rate increased with speed and for all seals mass-specific COT decreased with speed. Mass-specific metabolic rate was higher for subadult than adult fur seals and sea lions, corresponding to an overall higher minimum COT. Some sex differences were also apparent, such that female Australian fur seals and Australian sea lions had higher mass-specific metabolic rates than males. There were no species differences in standard or active metabolic rates for adult males or females. The seals in our study appear to operate at their physiological optimum during submerged swimming. However, the higher metabolic rates of young and female fur seals and sea lions may limit their scope for increasing foraging effort during times of resource limitation.

Searching for a truly “iso-metabolic” gas challenge in physiological MRI

Hypercapnia challenge (e.g. inhalation of CO2) has been used in calibrated fMRI as well as in the mapping of vascular reactivity in cerebrovascular diseases. An important assumption underlying these measurements is that CO2 is a pure vascular challenge but does not alter neural activity. However, recent reports have suggested that CO2 inhalation may suppress neural activity and brain metabolic rate. Therefore, the goal of this study is to propose and test a gas challenge that is truly “iso-metabolic,” by adding a hypoxic component to the hypercapnic challenge, since hypoxia has been shown to enhance cerebral metabolic rate of oxygen (CMRO2). Measurement of global CMRO2 under various gas challenge conditions revealed that, while hypercapnia (P = 0.002) and hypoxia (P = 0.002) individually altered CMRO2 (by −7.6 ± 1.7% and 16.7 ± 4.1%, respectively), inhalation of hypercapnic-hypoxia gas (5% CO2/13% O2) did not change brain metabolism (CMRO2 change: 1.5 ± 3.9%, P = 0.92). Moreover, cerebral blood flow response to the hypercapnic-hypoxia challenge (in terms of % change per mmHg CO2 change) was even greater than that to hypercapnia alone (P = 0.007). Findings in this study suggest that hypercapnic-hypoxia gas challenge may be a useful maneuver in physiological MRI as it preserves vasodilatory response yet does not alter brain metabolism.

Effects of pyrolytic and petrogenic polycyclic aromatic hydrocarbons on swimming and metabolic performance of zebrafish contaminated by ingestion

Depending on their origins, polycyclic aromatic hydrocarbons (PAH) are characterized by different chemical properties. Petrogenic PAH (e.g. from fossil fuels) and pyrolytic PAH (e.g. those produced by incineration processes) are therefore expected to affect organisms differently. The impact of trophic exposure to these PAH was investigated on swimming and metabolic performance of zebrafish Danio rerio. Two-month-old juveniles and six-month-old adults were individually challenged following a swimming step protocol. While pyrolytic exposure did not affect fish whatever the duration of exposure, it appeared that petrogenic PAH impaired adults' performance. Indeed, the active metabolic rate in petrogenic PAH-contaminated adults was significantly reduced by 35%, and critical swimming speed by 26.5%. This was associated with cardiac abnormalities, which are expected to contribute to the reduction of oxygen transport, particularly during intensive effort. These results may be due to the different composition and toxicity of PAH mixtures.

Acclimation temperature alters the relationship between growth and swimming performance among juvenile common carp (Cyprinus carpio)

Individual variation in growth, metabolism and swimming performance, their possible interrelationships, and the effects of temperature were investigated in 30 juvenile common carp (Cyprinus carpio) at two acclimation temperatures (15 and 25°C). We measured body mass, critical swimming speed (Ucrit), resting metabolic rate (RMR), active metabolic rate (AMR) and metabolic scope (MS) twice (28days apart) in both temperature groups. Fish acclimated to 25°C showed a 204% higher specific growth rate (SGR) than those acclimated to 15°C due to a 97% higher feeding rate (FR) and a 46% higher feed efficiency (FE). Among individuals, SGR was positively correlated with the FR and FE at both low and high temperatures. All measured variables (Ucrit, RMR and AMR) related to swimming except MS showed a high repeatability after adjusting for body mass (mass-independent). Fish acclimated to 25°C had a 40% higher Ucrit compared with 15°C acclimated fish, which was at least partially due to an improved metabolic capacity. AMR showed a 97% increase, and MS showed a 104% parallel increase with the higher acclimation temperature. Residual (mass-independent) Ucrit was positively correlated with residual RMR, AMR and MS, except for the residual RMR at high temperature. When acclimated to the lower temperature, both the residual and absolute Ucrit were negatively correlated with FR and FE and, hence, with SGR, suggesting a functional trade-off between growth and locomotion in fish acclimated to low temperatures. However, when acclimated to the higher temperature, this trade-off no longer existed; absolute Ucrit was positively correlated with SGR because individuals with rapid growth exhibited greatly increased body mass. The higher metabolic capacity at 25°C showed a positive effect on both swimming performance and growth rate (because of improved digestive efficiency) under the high-temperature condition, which we did not anticipate. Overall, these results indicate that temperature alters the relationship between growth and swimming performance of juvenile common carp. This change may be an adaptive strategy to seasonal temperature variation during their life history.

Metabolic rates of giant pandas inform conservation strategies.

The giant panda is an icon of conservation and survived a large-scale bamboo die off in the 1980s in China. Captive breeding programs have produced a large population in zoos and efforts continue to reintroduce those animals into the wild. However, we lack sufficient knowledge of their physiological ecology to determine requirements for survival now and in the face of climate change. We measured resting and active metabolic rates of giant pandas in order to determine if current bamboo resources were sufficient for adding additional animals to populations in natural reserves. Resting metabolic rates were somewhat below average for a panda sized mammal and active metabolic rates were in the normal range. Pandas do not have exceptionally low metabolic rates. Nevertheless, there is enough bamboo in natural reserves to support both natural populations and large numbers of reintroduced pandas. Bamboo will not be the limiting factor in successful reintroduction.

Associations between Resting, Activity, and Daily Metabolic Rate in Free-Living Endotherms: No Universal Rule in Birds and Mammals.

Energy management models provide theories and predictions for how animals manage their energy budgets within their energetic constraints, in terms of their resting metabolic rate (RMR) and daily energy expenditure (DEE). Thus, uncovering what associations exist between DEE and RMR is key to testing these models. Accordingly, there is considerable interest in the relationship between DEE and RMR at both inter- and intraspecific levels. Interpretation of the evidence for particular energy management models is enhanced by also considering the energy spent specifically on costly activities (activity energy expenditure [AEE] = DEE - RMR). However, to date there have been few intraspecific studies investigating such patterns. Our aim was to determine whether there is a generality of intraspecific relationships among RMR, DEE, and AEE using long-term data sets for bird and mammal species. For mammals, we use minimum heart rate (fH), mean fH, and activity fH as qualitative proxies for RMR, DEE, and AEE, respectively. For the birds, we take advantage of calibration equations to convert fH into rate of oxygen consumption in order to provide quantitative proxies for RMR, DEE, and AEE. For all 11 species, the DEE proxy was significantly positively correlated with the RMR proxy. There was also evidence of a significant positive correlation between AEE and RMR in all four mammal species but only in some of the bird species. Our results indicate there is no universal rule for birds and mammals governing the relationships among RMR, AEE, and DEE. Furthermore, they suggest that birds tend to have a different strategy for managing their energy budgets from those of mammals and that there are also differences in strategy between bird species. Future work in laboratory settings or highly controlled field settings can tease out the environmental and physiological processes contributing to variation in energy management strategies exhibited by different species.

The influence of carbon dioxide-induced water acidification on the osmotic and metabolic responses of the Baltic amphipod Gammarus oceanicus

The aim of this study was to investigate the effect of CO2-induced water acidification (pH values: 8.1 − control, 7.5 and 7.0) on the hemolymph osmolality and total metabolic rate of amphipod Gammarus oceanicus under short-term exposure. The hemolymph osmolality was measured using a vapor pressure osmometer. The metabolic rate was determined based on the heat dissipation measurements using a Calvet-type isothermal twin calorimeter. The hemolymph osmolality remained unchanged after exposure to pH 7.5 and it increased in the lowest treatment. Acidification did not significantly affect the resting or active metabolic rate. The observed results indicate that high osmolality maintained during hypercapnia did not generate additional physiological costs. It may therefore be concluded that the studied species is preadapted for this factor. The long-term adaptation to hypoosmotic conditions in the brackish Baltic Sea might also have preadapted it to hypercapnia because similar physiological mechanisms are responsible for the resistance to both factors.

Sexual dimorphism and physiological correlates of horn length in a South African isopod crustacean

Although isopod crustaceans frequently show sexual dimorphism in body size, sexual differences in other non-reproductive (secondary sexual) traits have been rarely studied. Furthermore, little is known about the physiological correlates of variation in the expression of sexually dimorphic traits in animals generally. Here, we show that the unusual dorsal horns of the South African horned isopod Deto echinata exhibit significantly different allometric scaling relationships with body size in males versus females. Not only are the horns of adult males substantially longer than those of adult females but also their log–log scaling slope with body length is significantly steeper than that of females and juveniles. Furthermore, in males relative horn length is positively correlated with body condition, as estimated by body mass relative to body length, and with resting metabolic (CO2 release) rate, also adjusted to body length. However, although showing a weak positive trend, the relative horn length of males is not significantly correlated with body length adjusted active metabolic rate. It is also not significantly correlated with the relative rates of whole body or cuticular water loss rates or with two measures of activity level. We hypothesize that the sexual dimorphism of horn length in D. echinata is the result of sexual selection and that horn length is a reliable indicator of the ability of a male to acquire and process resources, which in turn may be critical to female mate choice and (or) male–male competition for mates. However, this hypothesis requires further testing based on direct observations of female mate choice and male–male behavioral interactions, which have not yet been carried out.

Improving consumption rate estimates by incorporating wild activity into a bioenergetics model.

Consumption is the basis of metabolic and trophic ecology and is used to assess an animal's trophic impact. The contribution of activity to an animal's energy budget is an important parameter when estimating consumption, yet activity is usually measured in captive animals. Developments in telemetry have allowed the energetic costs of activity to be measured for wild animals; however, wild activity is seldom incorporated into estimates of consumption rates. We calculated the consumption rate of a free‐ranging marine predator (yellowtail kingfish, Seriola lalandi) by integrating the energetic cost of free‐ranging activity into a bioenergetics model. Accelerometry transmitters were used in conjunction with laboratory respirometry trials to estimate kingfish active metabolic rate in the wild. These field‐derived consumption rate estimates were compared with those estimated by two traditional bioenergetics methods. The first method derived routine swimming speed from fish morphology as an index of activity (a “morphometric” method), and the second considered activity as a fixed proportion of standard metabolic rate (a “physiological” method). The mean consumption rate for free‐ranging kingfish measured by accelerometry was 152 J·g−1·day−1, which lay between the estimates from the morphometric method (μ = 134 J·g−1·day−1) and the physiological method (μ = 181 J·g−1·day−1). Incorporating field‐derived activity values resulted in the smallest variance in log‐normally distributed consumption rates (σ = 0.31), compared with the morphometric (σ = 0.57) and physiological (σ = 0.78) methods. Incorporating field‐derived activity into bioenergetics models probably provided more realistic estimates of consumption rate compared with the traditional methods, which may further our understanding of trophic interactions that underpin ecosystem‐based fisheries management. The general methods used to estimate active metabolic rates of free‐ranging fish could be extended to examine ecological energetics and trophic interactions across aquatic and terrestrial ecosystems.

Respiration in spiders (Araneae).

Spiders (Araneae) are unique regarding their respiratory system: they are the only animal group that breathe simultaneously with lungs and tracheae. Looking at the physiology of respiration the existence of tracheae plays an important role in spiders with a well-developed tracheal system. Other factors as sex, life time, type of prey capture and the high ability to gain energy anaerobically influence the resting and the active metabolic rate intensely. Most spiders have metabolic rates that are much lower than expected from body mass; but especially those with two pairs of lungs. Males normally have higher resting rates than females; spiders that are less evolved and possess a cribellum have lower metabolic rates than higher evolved species. Freely hunting spiders show a higher energy turnover than spiders hunting with a web. Spiders that live longer than 1 year will have lower metabolic rates than those species that die after 1 year in which development and reproduction must be completed. Lower temperatures and starvation, which most spiders can cope with, will decrease the metabolic rate as well.

Trophic contamination by pyrolytic polycyclic aromatic hydrocarbons does not affect aerobic metabolic scope in zebrafish Danio rerio.

The effect of trophic exposure to pyrolitic polycyclic aromatic hydrocarbons (PAH) on aerobic metabolism of zebrafish Danio rerio was investigated. There were no significant differences in standard metabolic rate (SMR), active metabolic rate (AMR) or aerobic metabolic scope (AS) at any sublethal concentration of PAH in the diet of adult or juvenile fish. This suggests that under these experimental conditions, exposure to PAH in food did not influence aerobic metabolism of this species.

エネルギー代謝量からみた和船櫓漕ぎの推進効率の検討

The Japanese boat “Wasen” uses “Ro-Scull” as a human-powered propulsion system. These traditional boats and propulsion system are almost vanished in Japan due to decrease in material Wood, and boat-builder’s aging. “Ro-Scull” is thought to be effective propulsive method compared to other human-powered boat such as canoe and cutter boat. In this study, we evaluate propulsive efficiency of “Ro-Scull” based on active metabolic rate (MR). We measured MR and metabolic equivalents (Mets) during “Ro-Scull”, Walking, and jogging exercise by using indirect calorimeter. MR and Mets during “Ro-Scull” exercise were nearly equal to walking ones. Energy conversion efficiency of “Ro-scull” was 12.3%. These suggest that propulsion efficiency of “Ro-Scull” is efficient, and its exercise can be done for a prolonged time as well as Walking. “Ro-Scull” exercise seems to be useful way to become familiar with maritime activities and seamanship.

The Effect of CO2-Induced Seawater Acidification on the Behaviour and Metabolic Rate of the Baltic ClamMacoma balthica

The aim of this study was to investigate the effect of CO2-induced seawater acidification (pH 8.1; control pH 7.0 and 6.0) on the survival, behaviour and metabolic rate of the Baltic clam Macoma balthica during short- (36 hours) and long-term (14 days) exposures. Total metabolic rate and gaping behaviour (shell opening and closing) were determined based on heat dissipation measurements using an isothermal twin calorimeter of the Calvet type. The survival and burrowing behaviour during long-term exposure were observed every day. Short-term exposure to reduced pH did not significantly affect the resting or active metabolic rate. Neither were there any significant changes in gaping behaviour. However, long-term exposure significantly affected burrowing behaviour. In the lowest-pH treatment, both resting and active metabolic rates were also significantly higher than in the control.

Metabolism, oxidative stress and territorial behaviour in a female colour polymorphic cichlid fish

Intrasexual selection on body coloration is thought to play an important role in the evolution of colour polymorphism, but its physiological underpinnings have received limited attention. In the colour polymorphic cichlid Neochromis omnicaeruleus, three fully sympatric female colour morphs—a plain morph (P) and two conspicuously coloured blotched morphs, black-and-white blotched (WB) and orange blotched (OB)—differ in agonistic behaviour. We compared routine metabolic rate (when females were housed in social isolation), short-term energetic costs of interacting with a same-colour rival housed in an adjacent transparent chamber and oxidative stress between the three female colour morphs. WB females had a lower routine metabolic rate compared with the other colour morphs. WB females also had a lower active metabolic rate during inter-female interactions than OB females, while OB females used more oxygen per unit aggressive act than the other two colour morphs. However, there were no consistent differences in oxidative stress between the three morphs. Concerted divergence in colour, behaviour and metabolism might contribute to the evolution of these polymorphisms in sympatry.

Individual variation in metabolism and swimming performance in juvenile black carp (Mylopharyngodon piceus) and the effects of hypoxia

We investigated inter-individual variation in metabolism, swimming performance and the relationship between metabolism and swimming performance under normoxic and hypoxic oxygen conditions ([O2]) in juvenile black carp (Mylopharyngodon piceus). We measured the standard metabolic rate (SMR), critical swimming speed (Ucrit), active metabolic rate (AMR) and aerobic scope (AS) of 40 fish (body weight, 9.7–11.0 g; body length, 8.4–8.8 cm) at 20 °C under normoxic (100% air saturation) and hypoxic (30% air saturation) conditions. Hypoxia resulted in a significant decrease in all the investigated parameters (p < 0.05). The SMR, Ucrit and AMR exhibited consistent individual differences (repeatability) (p ≤ 0.022), whereas the AS had no consistency across different water [O2] conditions (p = 0.088). The SMR was positively correlated with the Ucrit, AMR and AS (p ≤ 0.002) under normoxic conditions. The SMR was also positively correlated with the Ucrit and AMR under hypoxic conditions (p ≤ 0.003) but there was no correlation between the SMR and AS (p = 0.141). The slope of the correlation between the SMR and Ucrit was shallower under hypoxic conditions than under normoxic conditions (F1,76 = 13.844, p < 0.001), which indicated that the swimming performance decreased more profoundly under hypoxic conditions in individuals with a high SMR.

Comparison of swimming capacity and energetics of migratory European eel (Anguilla anguilla) and New Zealand short-finned eel (A. australis).

The spawning migration of the European eel (Anguilla anguilla) can cover more than 6000 km, while that of the New Zealand short-finned eel (A. australis) is assumed to be approximately 3000 km. Since these species are expected to show adaptive traits to such an important lifetime event, we hypothesized differences in swimming capacity and energetics as a response to this adaptation. In an experimental swimming respirometer set-up, critical swimming speed (Ucrit), optimal swimming speed (Uopt), mass specific oxygen consumption rate (ṀO2), standard metabolic rate (SMR), active metabolic rate at Ucrit (AMRcrit) and at Uopt (AMRopt), the minimum cost of transport at Uopt (COTmin), and the scope for activity, were assessed and compared between the species. With a similar body length and mass, European eels showed ca. 25% higher values for both Ucrit and Uopt, and 23% lower values for COTmin, compared to New Zealand short-finned eels. However, SMR, AMRcrit, AMRopt, and scope for activity did not differ between the species, indicating very similar swimming physiology traits. This study discusses physiological aspects of long distance migration and provides recommendations for (a) swimming respirometry in anguilliform fish, and (b) telemetry research using externally attached pop-up tags.

Estimation of metabolic rate from activity measured by recorders deployed on Japanese sea bass Lateolabrax japonicus

Understanding the energy expenditure of top predators is important because a collapse in them could trigger trophic cascades through ecosystems. One such top predator, Japanese sea bass Lateolabrax japonicus, helps to balance the structure of the coastal marine ecosystem through predation. In this study, accelerometry was applied to the Japanese sea bass to estimate its energy expenditure under natural conditions. We attached accelerometers to five wild fish and measured metabolic rates such as the oxygen consumption rate (\({\dot{\text{V}}\text{O}}_{ 2}\), mg O2 kg−1 min−1) using a swim tunnel. Body beat frequency (BBF) was measured using the accelerometer. The BBF was correlated with the tail beat frequency (TBF) by analyzing video recordings. \({\dot{\text{V}}\text{O}}_{ 2}\) was related to swimming speed (U), TBF, and BBF. We estimated the standard (45.9 kJ kg−1 day−1) and active (124.0 kJ kg−1 day−1) metabolic rates when fish were not swimming and when they were swimming at the optimum swimming speed, respectively. The energy required to compensate for the above metabolic rates are between 83.3 and 275.6 kJ kg−1 day−1 using an assimilation efficiency of 0.7 and assuming that the growth rate is zero. These costs were comparable to consuming one or two prey fish per day (e.g., Japanese sardine: mean total length 155 ± SD 6 mm).

Modelling swimming activities and energetic costs in European sea bass (Dicentrarchus labrax L., 1758) during critical swimming tests

Muscular activity patterns in red and white muscles linked to oxygen consumption were studied during critical swimming tests in the sea bass (Dicentrarchus labrax Linnaeus 1758). The species is one of the most important for Mediterranean Sea aquaculture. A sigmoid model was used to fit both the oxygen consumption and red muscle activity while the white muscle activity pattern was described by an exponential model. Red muscle reaches an activation plateau close to critical swimming speed mostly due to the oxygen diffusion velocity in tissues. The exponential activation of white muscle appears to be linked to short and sudden periods of great energy need to cope with adverse conditions such as predation and escape. Both oxygen consumption and muscular activity were found to be size dependent. The bioenergetics of sea bass was modelled based on fish mass and swimming speed to predict the minimum and maximum speed as well as the mass-specific active metabolic rate and standard metabolic rate. An important finding was that contrary to other well-known species, swimming at subcritical speeds in sea bass involves both red and white muscle in different proportions.