Mechanical limits to maximum weapon size in a giant rhinoceros beetle

The horns of giant rhinoceros beetles are a classic example of the elaborate morphologies that can result from sexual selection. Theory predicts that sexual traits will evolve to be increasingly exaggerated until survival costs balance the reproductive benefits of further trait elaboration. In Trypoxylus dichotomus, long horns confer a competitive advantage to males, yet previous studies have found that they do not incur survival costs. It is therefore unlikely that horn size is limited by the theoretical cost–benefit equilibrium. However, males sometimes fight vigorously enough to break their horns, so mechanical limits may set an upper bound on horn size. Here, I tested this mechanical limit hypothesis by measuring safety factors across the full range of horn sizes. Safety factors were calculated as the ratio between the force required to break a horn and the maximum force exerted on a horn during a typical fight. I found that safety factors decrease with increasing horn length, indicating that the risk of breakage is indeed highest for the longest horns. Structural failure of oversized horns may therefore oppose the continued exaggeration of horn length driven by male–male competition and set a mechanical limit on the maximum size of rhinoceros beetle horns.     

Reduced oxygen at high altitude limits maximum size

The trend towards large size in marine animals with latitude, and the existence of giant marine species in polar regions have long been recognized, but remained enigmatic until a recent study showed it to be an effect of increased oxygen availability in sea water of a low temperature. The effect was apparent in data from 12 sites worldwide because of variations in water oxygen content controlled by differences in temperature and salinity. Another major physical factor affecting oxygen content in aquatic environments is reduced pressure at high altitude. Suitable data from high-altitude sites are very scarce. However, an exceptionally rich crustacean collection, which remains largely undescribed, was obtained by the British 1937 expedition from Lake Titicaca on the border between Peru and Bolivia in the Andes at an altitude of 3809 m. We show that in Lake Titicaca the maximum length of amphipods is 2-4 times smaller than other low-salinity sites (Caspian Sea and Lake Baikal).     

Theoretical limits to the correlation between pelagic larval duration and population genetic structure

Increasing dispersal duration should result in increasing dispersal distance, facilitating higher gene flow among populations. As such, it has long been predicted that genetic structure (e.g. F(ST) ) among populations of marine species should be strongly correlated with pelagic larval duration (PLD). However, previous studies have repeatedly shown a surprisingly poor correspondence. This result has been frequently interpreted as evidence for larval behaviours or physical oceanographic processes that result in larvae failing to reach their dispersal potential, or error inherent in estimating PLD and F(ST) . This study employed a computer modelling approach to explore the impacts of various uncertainties on the correlation between measures of genetic differentiation such as F(ST) and PLD. Results indicate that variation resulting from PLD estimation error had minor impacts on the correlation between genetic structure and PLD. However, variation in effective population size between species, errors in F(ST) estimation and non-equilibrium F(ST) values all had major impacts, resulting in dramatically weaker correlations between PLD and F(ST) . These results suggest that poor correlations between PLD and F(ST) may result from variation and uncertainty in the terms associated with the calculation of F(ST) values. As such, PLD may be a much stronger determinant of realized larval dispersal than suggested by the weak-to-moderate correlations between PLD and F(ST) reported in empirical studies.     

Theoretical analysis of rest and exercise hemodynamics in patients with total cavopulmonary connection

The objective of this study was to determine the impact of a total cavopulmonary connection on the main hemodynamic quantities, both at rest and during exercise, when compared with normal biventricular circulation. The analysis was performed by means of a mathematical model of the cardiovascular system. The model incorporates the main parameters of systemic and pulmonary circulation, the pulsating heart, and the action of arterial and cardiopulmonary baroreflex mechanisms. Furthermore, the effect of changes in intrathoracic pressure on venous return is also incorporated. Finally, the response to moderate dynamic exercise is simulated, including the effect of a central command, local metabolic vasodilation, and the "muscle pump" mechanism. Simulations of resting conditions indicate that the action of baroreflex regulatory mechanisms alone can only partially compensate for the absence of the right heart. Cardiac output and mean systemic arterial pressure at rest show a large decrease compared with the normal subject. More acceptable hemodynamic quantity values are obtained by combining the action of regulatory mechanisms with a chronic change in parameters affecting mean filling pressure. With such changes assumed, simulations of the response to moderate exercise show that univentricular circulation exhibits a poor capacity to increase cardiac output and to sustain aerobic metabolism, especially when the oxygen consumption rate is increased above 1.2-1.3 l/min. The model ascribes the poor response to exercise in these patients to the incapacity to sustain venous return caused by the high resistance to venous return and/or to exhaustion of volume compensation reserve.     

Ion permeability of the cytoplasmic membrane limits the maximum growth temperature of bacteria and archaea

Protons and sodium ions are the most commonly used coupling ions in energy transduction in bacteria and archaea. At their growth temperature, the permeability of the cytoplasmic membrane of thermophilic bacteria to protons is high compared with that of sodium ions. In some thermophiles, sodium is the sole energy-coupling ion. To test whether sodium is the preferred coupling ion at high temperatures, the proton- and sodium permeability was determined in liposomes prepared from lipids isolated from various bacterial and archaeal species that differ in their optimal growth temperature. The proton permeability increased with the temperature and was comparable for most species at their respective growth temperatures. Liposomes of thermophilic bacteria are an exception in the sense that the proton permeability is already high at the growth temperature. In all liposomes, the sodium permeability was lower than the proton permeability and increased with the temperature. The results suggest that the proton permeability of the cytoplasmic membrane is an important parameter in determining the maximum growth temperature.     

EMMLi: A maximum likelihood approach to the analysis of modularity

Identification of phenotypic modules, semiautonomous sets of highly correlated traits, can be accomplished through exploratory (e.g., cluster analysis) or confirmatory approaches (e.g., RV coefficient analysis). Although statistically more robust, confirmatory approaches are generally unable to compare across different model structures. For example, RV coefficient analysis finds support for both two‐ and six‐module models for the therian mammalian skull. Here, we present a maximum likelihood approach that takes into account model parameterization. We compare model log‐likelihoods of trait correlation matrices using the finite‐sample corrected Akaike Information Criterion, allowing for comparison of hypotheses across different model structures. Simulations varying model complexity and within‐ and between‐module contrast demonstrate that this method correctly identifies model structure and parameters across a wide range of conditions. We further analyzed a dataset of 3‐D data, consisting of 61 landmarks from 181 macaque (Macaca fuscata) skulls, distributed among five age categories, testing 31 models, including no modularity among the landmarks and various partitions of two, three, six, and eight modules. Our results clearly support a complex six‐module model, with separate within‐ and intermodule correlations. Furthermore, this model was selected for all five age categories, demonstrating that this complex pattern of integration in the macaque skull appears early and is highly conserved throughout postnatal ontogeny. Subsampling analyses demonstrate that this method is robust to relatively low sample sizes, as is commonly encountered in rare or extinct taxa. This new approach allows for the direct comparison of models with different parameterizations, providing an important tool for the analysis of modularity across diverse systems.     

A new approach to rowing ergometry: establishing exercise intensity relative to maximum force output

The present experiment evaluated a new approach to establish exercise intensity during hydraulic rowing ergometry. In contrast to the traditional approach where exercise intensity is augmented by systematically increasing workload, the new procedure increments the intensity of exercise while maintaining a constant percentage of maximum force output. Ten college females exercised on a hydraulic rower that allowed for control of rowing speed and resistance. The new method to establish work intensity was to row at a cadence of 30 c.min-1 at a force output equal to 50% of maximum rowing force at each setting determined dynamically prior to testing. Two protocols were used for the maximum tests on the hydraulic rower. Row 1 was a 17-min, six-stage, incremental continuous row test performed at increasingly difficult settings from easy (setting 1; 603 N) to difficult (setting 6; 893 N). Row 2 was identical to row 1 until 15 min when resistance was reduced to setting 2 (658 N) for "allout" effort during the last 2 min. During this time, cadence declined from 30 c.min-1 to 19.4 c.min-1 at dial setting 6 and increased to 35.4 c.min-1 at dial setting 2. Both rowing protocols were compared to maximal physiological responses during treadmill running (TM). Compared to TM, both rowing protocols elicited significantly lower maximum oxygen uptake (VO2max; P less than 0.05; row 1 = delta 29.0% and row 2 = delta 12.9%) and maximum heart rate (HRmax; P less than 0.05; row 1 = delta 12.9% and row 2 = delta 6.7%). Maximum ventilation (VEmax) during row 1 was also lower by 30.4% than TM (P less than 0.05).(ABSTRACT TRUNCATED AT 250 WORDS)     

Incubation capacity limits maximum clutch size in black-tailed gulls Larus crassirostris

Factors determining clutch size of birds have long been the central issue in studies in life histories. It is assumed that the configuration of brood patches could limit the maximum clutch size. To test this hypothesis we manipulated clutch sizes and measured egg temperature as well as reproductive consequences in black-tailed gulls Larus crassirostris , which usually lay two egg clutches and have three brood patches. Mean egg temperature in 4-egg clutches (32.6±1.0°C) was significantly lower than in 2-egg (34.6±0.4°C) and 3-egg clutches (34.1±0.4°C), because egg temperature of the coolest egg within a 4-egg clutch was often substantially lower than the other three eggs. The proportion of eggs hatching from 4-egg clutches (11.6%) was lower than those of 2-egg (49.1%) and 3-egg clutches (52.0%). Four-egg clutches had longer incubation periods (29.6±1.3 day) than 2-egg (28.1±1.7 day) and 3-egg clutches (28.0 ±1.3 day). The results indicate that incubation capacity, which may be determined by the configuration of brood patches, limits the maximum clutch size in black tailed gulls, but not the actual clutch size typically laid.     

Scaling of heat production by thermogenic flowers: limits to floral size and maximum rate of respiration

Effect of size of inflorescences, flowers and cones on maximum rate of heat production is analysed allometrically in 23 species of thermogenic plants having diverse structures and ranging between 1.8 and 600 g. Total respiration rate (, µmol s^?1) varies with spadix mass (M, g) according to in 15 species of Araceae. Thermal conductance (C, mW °C^?1) for spadices scales according to C = 18.5M^0.73. Mass does not significantly affect the difference between floral and air temperature. Aroids with exposed appendices with high surface area have high thermal conductance, consistent with the need to vaporize attractive scents. True flowers have significantly lower heat production and thermal conductance, because closed petals retain heat that benefits resident insects. The florets on aroid spadices, either within a floral chamber or spathe, have intermediate thermal conductance, consistent with mixed roles. Mass‐specific rates of respiration are variable between species, but reach 900 nmol s^?1 g^?1 in aroid male florets, exceeding rates of all other plants and even most animals. Maximum mass‐specific respiration appears to be limited by oxygen delivery through individual cells. Reducing mass‐specific respiration may be one selective influence on the evolution of large size of thermogenic flowers.     

Heavily loaded flight and limits to the maximum size of dragonflies (Anisoptera) and griffenflies (Meganisoptera)

An original hypothesis is presented that the maximum mass and size of living anisopteran dragonflies are constrained by a physiological performance limit: the wing muscle power required to permit reproductively successful males to carry heavier females in the so‐called ‘wheel position’ in flight. It is proposed that the same limit cannot have applied to all fossil Odonatoptera. As the physiology of the giant Carboniferous griffenfly Namurotypus sippeli precludes flight in the wheel position, it did not need to carry any substantial load aside from exogenous aerial prey. Based on its thorax dimensions, it is argued that Namurotypus flew with a relatively low maximum specific muscle power output in comparison with living Anisoptera. The extinction of some families of large Mesozoic Odonatoptera may have been exacerbated by competition with smaller (stem‐) Anisoptera that evolved higher specific power outputs and superior flight performance similar to living Anisoptera. To investigate the credibility of this flight‐performance size‐limit hypothesis and its consequences, an analysis of the scaling of the required flight power and available muscle power is presented using allometric relations. It is found that for living Anisoptera and fossil Odonatoptera, there are different limiting sizes, above which the required specific flight power would exceed the available muscle specific power. These limits are directly related to maximum load‐carrying capacity and the atmospheric air density at the habitual altitude. It is suggested that the largest living species of Petaluridae, Petalura ingentissima, is close to the proposed Anisoptera size limit at current near‐sea‐level air density conditions.     

Environmental and functional limits to muscular exercise and body size in marine invertebrate athletes

Many similarities exist between the key characteristics of muscular metabolism in marine invertebrates and those found in vertebrate striated muscle, even though there are important phosphagens and glycolytic end products that differ between groups. Lifestyles and modes of locomotion also vary extremely among invertebrates thereby shaping the pattern of exercise metabolism. In accordance with the limited availability of integrated ecological and physiological information the present paper reports recent progress in the exercise physiology of cephalopods, which are characterized by high rates of aerobic and anaerobic energy turnover during high velocity hunts or escapes in their pelagic environment, and a sipunculid worm, which mostly uses anaerobic resources during extended marathon-like digging excursions in the hypoxic marine sediment. Particular attention is paid to how lifestyle and oxygen availability in various marine environments shapes the use and rates of aerobic and anaerobic metabolism and acidosis as they depend on activity levels and energy saving strategies. Whereas aerobic scope and, accordingly, use of ambient oxygen by blood oxygen transport and skin respiration is maximized in some squids, aerobic scope is very small in the worm and anaerobic metabolism readily used upon muscular activity. Until recently, it was widely accepted that the glycolytic end product octopine, produced in the musculature of these invertebrates, acted as a weak acid and so did not compromise acid-base balance. However, it has now been demonstrated that octopine does cause acidosis. Concomitant study of tissue energy and acid-base status allows to evaluate the contribution of glycolysis, pH and free ADP accumulation to the use of the phosphagen and to the delayed drop in the Gibb's free energy change of ATP hydrolysis. The analysis reveals species specific capacities of these mechanisms to support exercise beyond the anaerobic threshold. During high intensity anaerobic exercise observed in squid, the levels of ATP free energy change finally fall to critical minimum levels contributing to fatigue. Maintenance of sufficiently high energy levels is found at low but extended rates of anaerobic metabolism as observed in the long term digging sipunculid worm. The greatest aerobic and anaerobic performance levels are seen in squid inhabiting the open ocean and appear to be made possible by the uniform and stable physicochemical parameters (esp. high O(2) and low CO(2) levels) that characterize such an environment. It is suggested that at least some squid operate at their functional and environmental limits. In extremely different environments, both the worm and the squids display a tradeoff between oxygen availability, temperature, performance level and also, body size.     

Genetic analysis of the maximum drinks phenotype

Using data provided by the Collaborative Study on the Genetics of Alcoholism we studied the genetics of a quantitative trait: the maximum number of drinks consumed in a 24-hour period. A two-stage method was used. First, linkage analysis was performed, followed by association analysis in regions where linkage was detected. Additionally, the extent of linkage disequilibrium among single-nucleotide polymorphisms (SNP) associated with the phenotype was assessed. Linkage to chromosomes 2 and 7 was detected, and follow-up association analysis found multiple trait-associated SNPs in the chromosome 7 linkage region. Chromosome 4, which has been implicated in previous studies of the maximum drinks phenotype, did not pass our threshold for linkage evidence in stage 1, but secondary analyses of this chromosome indicated modest evidence for both linkage and association. The evidence suggests that chromosome 7 may harbor an additional locus influencing the maximum drinks consumption phenotype.     

An acetyl group deficit limits mitochondrial ATP production at the onset of exercise

The oxygen deficit at the onset of submaximal exercise represents a period when the energy demand of contraction cannot be met solely by mitochondrial ATP generation, and as a consequence there is an acceleration of ATP re-synthesis from oxygen-independent routes (phosphocreatine hydrolysis and glycolysis). Historically, the origin of the oxygen deficit has been attributed to a lag in muscle blood flow and oxygen availability at the onset of exercise which limits mitochondrial respiration. However, more recent evidence suggests that considerable inertia exists at the level of mitochondrial enzyme activation and substrate supply. In support of this latter hypothesis, we have reported on a number of occasions that pharmacological activation of the pyruvate dehydrogenase complex (and consequent stockpiling of acetyl groups), using dichloroacetate or exercise interventions, can markedly reduce the degree of ATP re-synthesis from oxygen-independent routes during the rest-to-work transition period. This review will focus on these findings, and will offer the hypothesis that acetyl group delivery to the tricarboxylic acid cycle limits mitochondrial flux at the onset of exercise--the so-called acetyl group deficit.     

Max试验验证症状限制心肺运动试验为最大极限运动进一步临床研究*

目的: 验证临床受试者所完成的心肺运动试验（CPET）为最大极限运动,进一步设计完善Max试验验证CPET结果客观定量功能评估的准确性及以某特定指标的特定数值作为停止运动的标准是否可行。方法: 选择2017年9月至2019年1月在阜外医院签署知情同意书后进行CPET和Max试验受试者216例。其中正常受试者41例,因CPET峰值呼吸交换率（RER）≤1.10,或运动中心率和血压不上升,对CPET极限运动结果存在质疑的临床患者175例进行研究。其中60例已初步报告,本研究进一步扩大研究。Max试验方法：完成CPET测试后,先蹬车≥60 r/min,再施加130%峰值功率的恒定功率,鼓励受试者运动至不能坚持的极限状态。计算分析Max试验30 s的最大心率和最大摄氧量、及其与峰值心率和峰值摄氧量之间的差值和百分差值。百分差值=（Max值-峰值值）/Max值× 100%。评测标准：①若心率和摄氧量任一指标的差值百分比≤-10%（Max测试的数值低于CPET峰值数据）则定义Max试验操作失败,否则为成功;2若心率和摄氧量的差值百分比均在-10%~10%,则Max试验操作成功,证明CPET数据为极限运动,CPET 峰值相关数据较为准确;③若心率和摄氧量差值任一指标差值百分比≥10%时,则Max试验操作成功,证明CPET结果为非极限运动。结果: 病例组峰值摄氧量（L/min、ml/（min·kg）、%pred）、无氧阈（L/min、ml/（min·kg）、%pred）、峰值氧脉搏（ml/beat、%pred）、峰值RER、峰值收缩压（mmHg）、峰值运动负荷（W/min）、峰值心率（bpm）、摄氧有效性峰值平台（OUEP）（比值、%pred）低于正常组,二氧化碳通气有效性平均90 s最低值（Lowest Ve/VCO₂）（比值、%pred）、二氧化碳通气效率斜率（Ve/VCO₂ Slope）（比值、%pred）高于正常组（P<0.05）。所有正常组与病例组均安全无任何事件完成CPET和Max试验。216例受试者中,Max试验成功198例（91.7%）,其中证明CPET为极限运动182例,为非极限运动16例;失败18例（8.3%）。结论: 在临床检查中,若对CPET结果是否为最大极限存在质疑,利用Max试验可验证CPET是否为极限运动。Max试验方法安全可行,值得进一步深入研究和临床推广应用。