The energetic cost of mating in a promiscuous cephalopod

Costs that individuals incur through mating can play an important role in understanding the evolution of life histories and senescence, particularly in promiscuous species. Copulation costs, ranging from energy expenditure to reduced longevity, are widely studied in insects but have received substantially less attention in other taxa. One cost of mating, the energetic cost, is poorly studied across all taxa despite its potential importance for the many species where copulation is physically demanding and/or frequent. Here, we investigated the energetic cost of mating in both male and female dumpling squid (Euprymna tasmanica). In this species, copulation can last up to 3 h and requires that the male physically restrains the female. We report that the act of copulation halves the swimming endurance of both sexes, and that they take up to 30 min to recover. Such a reduction in post-copulatory performance may have important implications for predator avoidance, foraging ability and energy allocation. Therefore, quantifying this cost is essential to understand the evolution of reproductive strategies and behaviours such as female receptivity and male and female mating frequency.     

Effects of obesity and sex on the energetic cost and preferred speed of walking.

The metabolic energy cost of walking is determined, to a large degree, by body mass, but it is not clear how body composition and mass distribution influence this cost. We tested the hypothesis that walking would be most expensive for obese women compared with obese men and normal-weight women and men. Furthermore, we hypothesized that for all groups, preferred walking speed would correspond to the speed that minimized the gross energy cost per distance. We measured body composition, maximal oxygen consumption, and preferred walking speed of 39 (19 class II obese, 20 normal weight) women and men. We also measured oxygen consumption and carbon dioxide production while the subjects walked on a level treadmill at six speeds (0.50-1.75 m/s). Both obesity and sex affected the net metabolic rate (W/kg) of walking. Net metabolic rates of obese subjects were only approximately 10% greater (per kg) than for normal-weight subjects, and net metabolic rates for women were approximately 10% greater than for men. The increase in net metabolic rate at faster walking speeds was greatest in obese women compared with the other groups. Preferred walking speed was not different across groups (1.42 m/s) and was near the speed that minimized gross energy cost per distance. Surprisingly, mass distribution (thigh mass/body mass) was not related to net metabolic rate, but body composition (% fat) was (r2= 0.43). Detailed biomechanical studies of walking are needed to investigate whether obese individuals adopt novel energy saving mechanisms during walking.     

Muscle mechanical work requirements during normal walking: the energetic cost of raising the body's center-of-mass is significant

Inverted pendulum models of walking predict that little muscle work is required for the exchange of body potential and kinetic energy in single-limb support. External power during walking (product of the measured ground reaction force and body center-of-mass (COM) velocity) is often analyzed to deduce net work output or mechanical energetic cost by muscles. Based on external power analyses and inverted pendulum theory, it has been suggested that a primary mechanical energetic cost may be associated with the mechanical work required to redirect the COM motion at the step-to-step transition. However, these models do not capture the multi-muscle, multi-segmental properties of walking, co-excitation of muscles to coordinate segmental energetic flow, and simultaneous production of positive and negative muscle work. In this study, a muscle-actuated forward dynamic simulation of walking was used to assess whether: (1). potential and kinetic energy of the body are exchanged with little muscle work; (2). external mechanical power can estimate the mechanical energetic cost for muscles; and (3.) the net work output and the mechanical energetic cost for muscles occurs mostly in double support. We found that the net work output by muscles cannot be estimated from external power and was the highest when the COM moved upward in early single-limb support even though kinetic and potential energy were exchanged, and muscle mechanical (and most likely metabolic) energetic cost is dominated not only by the need to redirect the COM in double support but also by the need to raise the COM in single support.     

The energetic cost of maintaining lateral balance during human running

To quantify the energetic cost of maintaining lateral balance during human running, we provided external lateral stabilization (LS) while running with and without arm swing and measured changes in energetic cost and step width variability (indicator of lateral balance). We hypothesized that external LS would reduce energetic cost and step width variability of running (3.0 m/s), both with and without arm swing. We further hypothesized that the reduction in energetic cost and step width variability would be greater when running without arm swing compared with running with arm swing. We controlled for step width by having subjects run along a single line (zero target step width), which eliminated any interaction effects of step width and arm swing. We implemented a repeated-measures ANOVA with two within-subjects fixed factors (external LS and arm swing) to evaluate main and interaction effects. When provided with external LS (main effect), subjects reduced net metabolic power by 2.0% (P = 0.032) and step width variability by 12.3% (P = 0.005). Eliminating arm swing (main effect) increased net metabolic power by 7.6% (P < 0.001) but did not change step width variability (P = 0.975). We did not detect a significant interaction effect between external LS and arm swing. Thus, when comparing conditions of running with or without arm swing, external LS resulted in a similar reduction in net metabolic power and step width variability. We infer that the 2% reduction in the net energetic cost of running with external LS reflects the energetic cost of maintaining lateral balance. Furthermore, while eliminating arm swing increased the energetic cost of running overall, arm swing does not appear to assist with lateral balance. Our data suggest that humans use step width adjustments as the primary mechanism to maintain lateral balance during running.     

The energetic cost of locomotion: humans and primates compared to generalized endotherms

A wide range of selective pressures have been advanced as possible causes for the adoption of bipedalism in the hominin lineage. One suggestion has been that because modern human walking is relatively efficient compared to that of a typical quadruped, the ancestral quadruped may have reaped an energetic advantage when it walked on two legs. While it has become clear that human walking is relatively efficient and human running inefficient compared to "generalized endotherms", workers differ in their opinion of how the cost of human bipedal locomotion compares to that of a generalized primate walking quadrupedally. One view is that human walking is particularly efficient in comparison to other primates. The present study addresses this by comparing the cost of human walking and running to that of the eight primate species for which data are available and by comparing cost in primates to that of a "generalized endotherm". There is no evidence that primate locomotion is more costly than that of a generalized endotherm, although more data on adult Old World monkeys and apes would be useful. Further, human locomotion does not appear to be particularly efficient relative to that of other primates.     

Buccal cell DNA extraction: yield, purity, and cost: a comparison of two methods

Simple and cost-effective methods are needed to extract DNA in order to use it in large-scale studies. Blood is an excellent DNA source; however, it is costly and invasive thus an alternative is needed. Several kits and chemical protocols using buccal cells have been proposed for DNA extraction. The objective of the study is to evaluate buccal NaOH chemical protocol and Nucleospin Tissue Kit (BD Biosciences, Macery-Nagel, Germany) for DNA extraction. DNA swab samples were collected from 300 voluntary participants. DNA yields and purity were measured by NaOH and Nucleospin Tissue Kit techniques; the cost and time consumption for DNA extraction per sample were assessed as well. Results have shown that DNA amount and purity extracted by NaOH procedure was compared to that of the kit (p = 0.164; p = 0.249, respectively). NaOH method was considered cheaper and less time consuming (0.06 versus 3.80 USD, and 1.33 versus 3.59 minutes per sample, p < 0.001). Buccal cell derived DNA extracted by NaOH protocol can be considered a feasible substitute for more expensive and time-consuming kits.     

The energy cost of walking or running on sand

Oxygen uptake (VO2) at steady state, heart rate and perceived exertion were determined on nine subjects (six men and three women) while walking (3-7 km.h-1) or running (7-14 km.h-1) on sand or on a firm surface. The women performed the walking tests only. The energy cost of locomotion per unit of distance (C) was then calculated from the ratio of VO2 to speed and expressed in J.kg-1.m-1 assuming an energy equivalent of 20.9 J.ml O2-1. At the highest speeds C was adjusted for the measured lactate contribution (which ranged from approximately 2% to approximately 11% of the total). It was found that, when walking on sand, C increased linearly with speed from 3.1 J.kg-1.m-1 at 3 km.h-1 to 5.5 J.kg-1.m-1 at 7 km.h-1, whereas on a firm surface C attained a minimum of 2.3 J.kg-1.m-1 at 4.5 km.h-1 being greater at lower or higher speeds. On average, when walking at speeds greater than 3 km.h-1, C was about 1.8 times greater on sand than on compact terrain. When running on sand C was approximately independent of the speed, amounting to 5.3 J.kg-1.m-1, i.e. about 1.2 times greater than on compact terrain. These findings could be attributed to a reduced recovery of potential and kinetic energy at each stride when walking on sand (approximately 45% to be compared to approximately 65% on a firm surface) and to a reduced recovery of elastic energy when running on sand.     

Predicting metabolic cost of level walking

Energy expenditure in walking is usually expressed as a function of walking speed. However, this relationship applies only to freely adopted step length-step rate patterns. Both the step length and the step rate must be used to preduct the energy expenditure for any combination of step length and step rate. Evidence on seven subjects indicates that the energy demand for such a combination can be determined by conducting two experiments. In the first, the subject is allowed to freely choose his own walking pattern to achieve a set of prescribed speeds. In the second, the speed is kept constant but the subject is forced to adopt a range of prescribed step rates. The results of the two experiments combined yield enough data to make possible the determination of the energy equation of the pattern, encompassing both "free" and "forced" gaits. Results show that the freely chosen step rate requires the least oxygen consumption at any given speed. Any other forced step rate at the same speed increases the oxygen cost over that required for the "free" step rate.     

The energetic cost of reproductive conflicts in the ant Pachycondyla obscuricornis

In a variety of social animals, individuals can secure reproductive rights through aggressive dominance. Direct individual benefits of aggression are widely recognized, but underlying costs affecting group productivity, and thus indirect benefits, are less clear. Costs of aggressive regulation of reproduction are especially important in small social insect colonies, where individual workers could potentially dominate male production. We estimated the energetic costs associated with the regulation of worker reproduction in the ponerine ant Pachycondyla obscuricornis, using the total CO2 emission of a colony as a measure. The level of CO2 emission of 12 experimental colonies varied significantly during five periods with varying levels of aggression and egg-laying. Overall, CO2 emission increased with the degree of fighting in a colony, but was not associated with differences in egg-laying. Aggressive regulation of reproduction and the formation of a dominance hierarchy thus pose an energetic cost to the colony. Furthermore, workers reduce their work-activities immediately after experimental orphaning, giving a further cost to the colony. These costs might influence the outcome of conflicts over male production in ants. This paper presents the first quantification of energetic costs of aggressive behavior regulating reproduction in ants.     

The cost of sex: quantifying energetic investment in gamete production by males and females

The relative energetic investment in reproduction between the sexes forms the basis of sexual selection and life history theories in evolutionary biology. It is often assumed that males invest considerably less in gametes than females, but quantifying the energetic cost of gamete production in both sexes has remained a difficult challenge. For a broad diversity of species (invertebrates, reptiles, amphibians, fishes, birds, and mammals), we compared the cost of gamete production between the sexes in terms of the investment in gonad tissue and the rate of gamete biomass production. Investment in gonad biomass was nearly proportional to body mass in both sexes, but gamete biomass production rate was approximately two to four orders of magnitude higher in females. In both males and females, gamete biomass production rate increased with organism mass as a power law, much like individual metabolic rate. This suggests that whole-organism energetics may act as a primary constraint on gamete production among species. Residual variation in sperm production rate was positively correlated with relative testes size. Together, these results suggest that understanding the heterogeneity in rates of gamete production among species requires joint consideration of the effects of gonad mass and metabolism.     

Swimming by sea otters: adaptations for low energetic cost locomotion

The energetics and hydrodynamics of surface and submerged swimming were compared in the sea otter (Enhydra lutris). 1. Sea otters used two distinct speed ranges that varied with swimming mode. Sustained surface swimming was limited to speeds less than 0.80 m/s, while sustained submerged swimming occurred over the range of 0.60 to 1.39 m/s. 2. Rates of oxygen consumption (VO2) at the transition speed (0.80 m/s) were 41% lower for submerged swimming by sea otters in comparison to surface swimming. 3. Total cost of transport for surface swimming sea otters, 12.56 joules/kg.m, was more than 12 times the predicted value for a similarly-sized salmonid fish. Transport costs for submerged swimming at the same speed was only 7.33 times the predicted value. 4. The allometric relationship for minimum cost of transport in surface swimming birds and mammals was y = 23.87 chi -0.15 where y = cost of transport in joules/kg.m and x = body mass in kg. This regression loosely parallels the relationship for salmonid fish. 5. Correlations between aquatic behavior, morphological specialization, and swimming energetics indicate that the development of swimming in mustelids involved transitions from fore-paw to hind-paw propulsion, and from surface to submerged swimming.     

The relative cost of bent-hip bent-knee walking is reduced in water

The debate about how early hominids walked may be characterised as two competing hypotheses: They moved with a fully upright (FU) gait, like modern humans, or with a bent-hip, bent-knee (BK) gait, like apes. Both have assumed that this bipedalism was almost exclusively on land, in trees or a combination of the two. Recent findings favoured the FU hypothesis by showing that the BK gait is 50–60% more energetically costly than a FU human gait on land. We confirm these findings but show that in water this cost differential is markedly reduced, especially in deeper water, at slower speeds and with greater knee flexion. These data suggest that the controversy about australopithecine locomotion may be eased if it is assumed that wading was a component of their locomotor repertoire and supports the idea that shallow water might have been an environment favourable to the evolution of early forms of “non-optimal” hominid bipedalism.     

The energetic cost of the fever response in three species of ectothermic vertebrates

Three species of ectothermic vertebrates: goldfish (Carassius auratus), green tree frogs (Hyla cinerea), and desert iguanas (Dipsosaurus dorsalis) were used in this study. Metabolic rates for each species were determined at the normal afebrile preferred body temperature and at the febrile preferred body temperature or other higher body temperatures. The febrile metabolic rate (or higher rate) was significantly greater than the afebrile metabolic rate (or lower rate) in each species. The average increase in energetic cost for goldfish and desert iguana was 64.5% while the increase for the green tree frog was between 24 and 70%.     

Predicting the metabolic cost of incline walking from muscle activity and walking mechanics

The goal of this study was to identify which muscle activation patterns and gait features best predict the metabolic cost of inclined walking. We measured muscle activation patterns, joint kinematics and kinetics, and metabolic cost in sixteen subjects during treadmill walking at inclines of 0%, 5%, and 10%. Multivariate regression models were developed to predict the net metabolic cost from selected groups of the measured variables. A linear regression model including incline and the squared integrated electromyographic signals of the soleus and vastus lateralis explained 96% of the variance in metabolic cost, suggesting that the activation patterns of these large muscles have a high predictive value for metabolic cost. A regression model including only the peak knee flexion angle during stance phase, peak knee extension moment, peak ankle plantarflexion moment, and peak hip flexion moment explained 89% of the variance in metabolic cost; this finding indicates that kinematics and kinetics alone can predict metabolic cost during incline walking. The ability of these models to predict metabolic cost from muscle activation patterns and gait features points the way toward future work aimed at predicting metabolic cost when gait is altered by changes in neuromuscular control or the use of an assistive technology.     

Disrupting Escherichia coli: a comparison of methods

The often-encountered problem of disrupting bacteria for the purpose of extracting soluble protein has generated various methods. Many require specialized equipment. Very often, especially during preliminary studies, investigators need a simple, fast, and inexpensive method for cell disruption that preserves biological activity. This paper compares some simple and inexpensive methods for cell disruption, such as bead-vortexing, freezing-thawing, French pressing, and sonication. It also provides some tips to increase protein yield and preserve biological activity. If performed under optimal conditions, bead-vortexing gives protein yields that are comparable to French pressing and sonication. It also preserves the activities of labile enzymes and releases periplasmic enzymes. Vortexing with glass beads appears to be the simplest method for cell disruption.     

Gonorrhea modeling: a comparison of control methods

A population dynamics model for a heterogeneous population is used to compare the effectiveness of six prevention methods for gonorrhea involving population screening and contact tracing of selected groups. The population is subdivided according to sex, sexual activity, and symptomatic or asymptomatic infection. For this model contact tracing of certain groups is more effective than general population screening.