Energetic Hypothesis Research Articles

Disentangling Positive Selection from Relaxed Selection in Animal Mitochondrial Genomes.

AbstractDisentangling different types of selection is a common goal in molecular evolution. Elevated dN/dS ratios (the ratio of nonsynonymous to synonymous substitution rates) in focal lineages are often interpreted as signs of positive selection. Paradoxically, relaxed purifying selection can also result in elevated dN/dS ratios, but tests to distinguish these two causes are seldomly implemented. Here, we reevaluated seven case studies describing elevated dN/dS ratios in animal mitochondrial DNA (mtDNA) and their accompanying hypotheses regarding selection. They included flightless lineages versus flighted lineages in birds, bats, and insects and physiological adaptations in snakes, two groups of electric fishes, and primates. We found that elevated dN/dS ratios were often not caused by the predicted mechanism, and we sometimes found strong support for the opposite mechanism. We discuss reasons why energetic hypotheses may be confounded by other selective forces acting on mtDNA and caution against overinterpreting singular molecular signals, including elevated dN/dS ratios.

A day at the beach: Does visually perceived distance depend on the energetic cost of walking?

It takes less effort to walk from here to the Tiki Hut on the brick walkway than on the sandy beach. Does that influence how far away the Tiki Hut looks? The energetic cost of walking on dry sand is twice that of walking on firm ground (Lejeune et al., 1998). If perceived distance depends on the energetic cost or anticipated effort of walking (Proffitt, 2006), then the distance of a target viewed over sand should appear much greater than one viewed over brick. If perceived distance is specified by optical information (e.g., declination angle from the horizon; Ooi et al., 2001), then the distances should appear similar. Participants (N = 13) viewed a target at a distance of 5, 7, 9, or 11 m over sand or brick and then blind-walked an equivalent distance on the same or different terrain. First, we observed no main effect of walked terrain; walked distances on sand and brick were the same (p = 0.46), indicating that locomotion was calibrated to each substrate. Second, responses were actually greater after viewing over brick than over sand (p < 0.001), opposite to the prediction of the energetic hypothesis. This unexpected overshooting can be explained by the slight incline of the brick walkway, which partially raises the visually perceived eye level (VPEL) and increases the target distance specified by the declination angle. The result is thus consistent with the information hypothesis. We conclude that visually perceived egocentric distance depends on optical information and not on the anticipated energetic cost of walking.

High ATP Production Fuels Cancer Drug Resistance and Metastasis: Implications for Mitochondrial ATP Depletion Therapy.

Recently, we presented evidence that high mitochondrial ATP production is a new therapeutic target for cancer treatment. Using ATP as a biomarker, we isolated the “metabolically fittest” cancer cells from the total cell population. Importantly, ATP-high cancer cells were phenotypically the most aggressive, with enhanced stem-like properties, showing multi-drug resistance and an increased capacity for cell migration, invasion and spontaneous metastasis. In support of these observations, ATP-high cells demonstrated the up-regulation of both mitochondrial proteins and other protein biomarkers, specifically associated with stemness and metastasis. Therefore, we propose that the “energetically fittest” cancer cells would be better able to resist the selection pressure provided by i) a hostile micro-environment and/or ii) conventional chemotherapy, allowing them to be naturally-selected for survival, based on their high ATP content, ultimately driving tumor recurrence and distant metastasis. In accordance with this energetic hypothesis, ATP-high MDA-MB-231 breast cancer cells showed a dramatic increase in their ability to metastasize in a pre-clinical model in vivo. Conversely, metastasis was largely prevented by treatment with an FDA-approved drug (Bedaquiline), which binds to and inhibits the mitochondrial ATP-synthase, leading to ATP depletion. Clinically, these new therapeutic approaches could have important implications for preventing treatment failure and avoiding cancer cell dormancy, by employing ATP-depletion therapy, to target even the fittest cancer cells.

A Day at the Beach: Does the energy cost of walking influence visually perceived distance?

It takes less effort to walk from here to the Tiki Hut on the brick walkway than on the sandy beach. Does that influence how far away the Tiki Hut looks? Last year at VSS we ran the experiment to find out. The energy cost of walking on dry sand is twice that of walking on firm ground (Lejeune, et al., 1998). If distance is perceived in units of energetic cost or ‘behavioural potential’ (Proffitt, 2006), then visual distances over sand should be double those over brick. If distance is perceived in constant units (e.g. eye-heights as specified by declination angle; Ooi, et al., 2001), then the visual distances should be similar. Participants (N=13) viewed a target at a distance of 5, 7, 9, or 11m over sand or brick, were blindfolded and turned toward the same or different terrain, then blind-walked to match the visual distance. Walked Distance was analyzed using mixed effects regression with Target Distance, Viewed Terrain, and Walked Terrain as fixed-effect predictors. The energetic hypothesis predicts that walked distance should be greater after viewing over sand than over brick. But walked distance was actually shorter after viewing over sand (β = -0.59, SE = 0.21, χ2(1) = 6.05, p = 0.01). There was no difference between walked distance on sand and brick (p = 0.71), with moderate evidence for the null hypothesis (BFJZS = 3.38). Thus, perceived distance did not increase with the energy cost of the viewed terrain. Moreover, walked distance was no different on sand and brick, indicating an accurate locomotor calibration to each terrain. The results imply that distance is not perceived in terms of energetic cost. Rather, visually perceived distance is constant, and walking is guided by a visual-locomotor mapping (e.g. from declination angle to locomotor distance on the terrain) (Warren, 2019).

Two Processes Regulating Trophic Energy Flow in Pelagic and Terrestrial Ecosystems: Trophic Efficiency and Body Size–Dependent Biomass Production: (A Reply to Giacomini)

AbstractWe proposed two hypotheses to explain why food chains are longer in pelagic than terrestrial ecosystems: greater trophic efficiency of pelagic animal taxa at lower trophic levels and a higher pelagic biomass production rate at lower trophic levels because of smaller pelagic body masses. Giacomini favored the former, invoking in support the energetic equivalence hypothesis. We reply that the energetic equivalence hypothesis does not describe populations at differing trophic levels and so does not refute a significant role for body-mass dependence in explaining faster trophic transfer in pelagic ecosystems. Metabolic scaling as body mass to the exponent 1/4, widely accepted, remains important for trophic dynamic models. We suggest a likelihood method to compare the two hypotheses on the basis of models of whole-ecosystem energetics.

Counterpoint

Claims about alterations in perception based on manipulations of the energetics hypothesis (and other influences) are often framed as interesting specifically because they affect our perceptual experience. Many control experiments conducted on such perceptual effects suggest, however, that they are the result of attribution effects and other kinds of judgmental biases influencing the reporting process rather than perception itself. Schnall (2017, this issue), appealing to Heider’s work on attribution, argues that it is fruitless to try to distinguish between perception and attribution. This makes the energetics hypothesis less interesting.

THE FORMATION OF THE SURFACE DURING AN ABRASIVE FINISHING AT CONSTANT AND VARIABLE CLAMPING FORCES: FINISHING AT CONSTANT AND VARIABLE CLAMPING FORCES

In this work the flat abrasive finishing process is reported. The mechanical and chemical phenomena taking place during the finishing process are described. The most common forming process schemes observed during abrasive treatment are reviewed. The schemes of pulsed, geometrical and energetic models are discussed. On the basis of the energetic hypothesis and Preston hypothesis, the rational law of clamping force variation during the finishing is established. This law allowed for the stabilization of the contact pressure and an increase of the treatment efficiency. An exponential dependence of a boundary contact area variation during the finishing of planes with different initial profiles of the macro-topography is developed. An experimental confirmation of the Preston hypothesis during the abrasive finishing is carried out with the plane-finishing machine “Rastr 220”. The comparison experiments are carried out at constant and variable clamping forces of the treated surface towards the tool. It is found that by varying the forces according to the exponential law (i.e. similar to the changing of the boundary contact area during the abrasive finishing) the process efficiency is increasing by a factor of 2.5–3. It is shown that the changes of the root mean square deviation [Formula: see text] of the roughness [Formula: see text] of a treated surface in the beginning of the process (at time [Formula: see text] min) during the finishing at a constant clamping force is two times higher compared to the treatment at a variable clamping force in the same time period.

Calcium carbonate saturation state: on myths and this or that stories

Abstract In a recent opinion article titled “The Omega Myth”, Cyronak et al. provide a series of arguments as to why saturation state should not matter to marine calcifiers. In sections of their article, they highlight several aspects of our published work, and unfortunately appear to misinterpret the foundation for the kinetic–energetic hypothesis we have laid out previously. While we are in full agreement that omega sensitivity is not a substrate limitation issue, we more clearly detail below what a kinetic limitation means and why it is different from a substrate limitation. The kinetic argument we have previously presented highlights how the energetic cost of calcification increases with a decreasing saturation state (or omega). We then highlight several issues with a bicarbonate/proton flux model applied to newly developing marine bivalve larvae, and discuss how a bicarbonate/proton flux and omega-based sensitivity model do not have to be mutually exclusive. Our intent with this comment is to clarify the points raised by Cyronak et al. about our work, and help to move the thinking past dialectic debate towards a more synthetic view on ocean acidification impacts on marine calcifiers.

Size-Energy Relationships in Ecological Communities

Hypotheses that relate body size to energy use are of particular interest in community ecology and macroecology because of their potential to facilitate quantitative predictions about species interactions and to clarify complex ecological patterns. One prominent size-energy hypothesis, the energetic equivalence hypothesis, proposes that energy use from shared, limiting resources by populations or size classes of foragers will be independent of body size. Alternative hypotheses propose that energy use will increase with body size, decrease with body size, or peak at an intermediate body size. Despite extensive study, however, size-energy hypotheses remain controversial, due to a lack of directly-measured data on energy use, a tendency to confound distinct scaling relationships, and insufficient attention to the ecological contexts in which predicted relationships are likely to occur. Our goal, therefore, was to directly evaluate size-energy hypotheses while clarifying how results would differ with alternate methods and assumptions. We comprehensively tested size-energy hypotheses in a vertebrate frugivore guild in a tropical forest in Madagascar. Our test of size-energy hypotheses, which is the first to examine energy intake directly, was consistent with the energetic equivalence hypothesis. This finding corresponds with predictions of metabolic theory and models of energy distribution in ecological communities, which imply that body size does not confer an advantage in competition for energy among populations or size classes of foragers. This result was robust to different assumptions about energy regulation. Our results from direct energy measurement, however, contrasted with those obtained with conventional methods of indirect inference from size-density relationships, suggesting that size-density relationships do not provide an appropriate proxy for size-energy relationships as has commonly been assumed. Our research also provides insights into mechanisms underlying local size-energy relationships and has important implications for predicting species interactions and for understanding the structure and dynamics of ecological communities.

Sea urchin embryos exposed to cadmium as an experimental model for studying the relationship between autophagy and apoptosis

The sea urchin embryo is a suitable model that offers an excellent opportunity to investigate different defence strategies activated in stress conditions. We previously showed that cadmium accumulates in a dose- and time-dependent manner into embryonic cells, activating different stress and defence mechanisms, including the synthesis of HSPs and the onset of apoptosis and/or autophagy. In this paper we investigated the functional relationship between autophagy and apoptosis, evaluating apoptosis signals in cadmium-exposed Paracentrotus lividus embryos with inhibited autophagy. We found that the inhibition of autophagy produced the concurrent reduction of apoptosis, suggesting that the two phenomena are functionally related. Considering the catabolic role of autophagy, an energetic hypothesis to explain the relationship was evaluated. Using a substrate for ATP production, we found that apoptosis, assessed by TUNEL and cleaved caspase-3 immunocytochemistry, was substantially restored in cadmium-treated embryos where autophagy was inhibited by 3-Methyladenine. On the basis of these results, we propose that, autophagy could play a crucial role in stress response of this organism because autophagy could energetically contribute to apoptotic execution through its catabolic role.

Reproductive parameters of female orangutans (Pongo pygmaeus wurmbii) 1971–2011, a 40-year study at Tanjung Puting National Park, Central Kalimantan, Indonesia

This study presents reproductive parameter data gathered by direct observation over a 40-year period (1971-2011) of the provisioned free-ranging population of orangutans at Camp Leakey in Tanjung Puting National Park, Central Kalimantan, Indonesia. Age at first reproduction, interbirth interval (IBI), sex ratio at birth, and infant mortality for 19 female orangutans (11 first-generation wild-born ex-captive mothers and 8 second-generation mothers) are included in this analysis. Age at first reproduction among the first-generation mothers was similar to that among wild orangutans, while second-generation mothers had a significantly earlier age at first reproduction. IBIs were similar among first- and second-generation mothers and were significantly shorter than those recorded in studies of wild orangutan populations. There was an expected male-biased sex ratio at birth and a slightly higher than expected rate of infant mortality when compared to wild populations. Infant mortality was primarily seen among second-generation mothers who gave birth before the age of 12, and among first births of some first-generation mothers. These results lend support to the ecological energetics hypothesis, which predicts that increased diet quality leads to a faster rate of reproduction.

Cheating for sex: inherent disadvantage or energetic constraint?

In many species, some males gain access to females by cheating, i.e. by exploiting the reproductive investment made by other males. There are two main nonmutually exclusive hypotheses to explain the decision of an individual to switch from the classical reproductive tactic to the parasitic one: the inherent disadvantage and energetic constraint hypotheses. For the first time, we tested and compared these two hypotheses in an experimental design by manipulating food availability of males with different degrees of inherent disadvantage in the caller/satellite system of Hyla arborea. Small males were more likely to be satellites than large individuals as predicted by the inherent disadvantage hypothesis. However, food deprived males were not more likely to be satellites than those that were fed, suggesting that the energetic constraint is not the main process explaining the use of a parasitic reproductive tactic. Moreover, all else being equal, males were more likely to act as satellites if confronted with an attractive competitor than if confronted with an unattractive one. In addition, males had a greater probability of being satellites at the beginning and end of the night, whereas they were more likely to call during the middle of the night. These results could highlight synchronization between male tactics and expected female availability during the course of the night.

Body size distributions in North American freshwater fish: large‐scale factors

ABSTRACTAim To document continental‐ and regional‐scale variation in the size distributions of freshwater fish and examine some energetic, evolutionary and biogeographic explanations for these patterns.Location North America.Methods Regional species lists, coupled with habitat and body size information, were used to document the spatial patterns.Results At the continental scale, riverine specialist fishes show a unimodal, right‐skewed, body size distribution whereas habitat generalist and lacustrine specialist species exhibit bimodal size distributions, with only a slight preponderance of small‐mode species. Most large‐mode species are migratory. Resident species, unlike migratory ones, show a latitudinal increase in mean size, but the size increase across all species is steeper because the importance of large migratory species increases with latitude. Size distributions change from right‐ to left‐skewed with increasing latitude. Maximum body size does not change with increasing family richness but minimum size declines and skewness increases, consistent with diversification of small species. Skewness does not vary with mean family body size.Main conclusions Post‐glacial recolonization by large, habitat generalist, migratory species is the main determinant of latitudinal size distribution trends. There is little support for the energetic hypothesis, but the data are consistent with a negative Cope's rule.

Arrival timing in subadult and adult Black Redstart males: competition-dependent behaviour?

The different arrival times of 1-year-old and older males is a widely recognised phenomenon in most migrating passerines. The converse pattern, i.e. the yearlings arriving at the breeding grounds at the same time as adults, has been reported only exceptionally. Two hypotheses have been formulated to explain the delayed arrival of yearling males: investment reduction, and energetic constraint hypotheses, respectively. However, which of these is the more appropriate to explain the phenomenon remains unclear. We studied arrival timing in two distinct urban populations of Black Redstart (Phoenicurus ochruros), a small passerine exhibiting delayed plumage maturation, whereby dull-coloured yearlings are easily distinguishable from black adults. The birds inhabiting South Moravian villages did not show differential timing of arrivals between yearlings and older birds in the resource-rich environment. On the other hand, under suboptimal conditions in Prague the yearlings were delayed by three days on average compared to the adult males. We discuss the possibility that the young birds of this species sample the situation on a potential breeding site during the autumnal singing period and flexibly adapt their migrating behaviour according to the circumstances. In such a case the investment reduction hypothesis is the best explanation for different arrival timing of yearling and adult males in Black Redstart.

Diversity, Body Mass, and Latitudinal Gradients in Primates

We examined latitudinal gradients in central value and diversity of body mass of primates to increase understanding of the Forster effect (decrease in taxonomic diversity with increasing latitude) and the Bergmann effect (increase of body mass with latitude). Data are from the literature. We used species’ median body mass of females and mid-latitude (N = 164). We account for phylogenetic effects with comparative analysis by independent contrasts and analysis at differing taxonomic levels. Globally, diversity of both taxonomy and body mass declined significantly with increasing latitude. The decrease in the range of body masses with increasing latitude was caused mainly by the absence of several small-bodied clades at higher latitudes, but also by the absence there of great apes. The disappearance of the small-bodied clades caused an increase in median body mass with latitude, i.e., primates show a significant taxon-wide Bergmann effect, including with phylogenetic correction. Within the Primates, the Bergmann effect was significant within taxa that extend from the equator the farthest into temperate regions: the Old World infraorder Catarrhini, family Cercopithecidae, and subfamily Cercopithecinae; the Asian Cercopithecidae; and in Southeast Asian Sunda, Macaca. The results accord with hypotheses for the Forster effect that latitudinal gradients in taxonomic diversity result from high rates of speciation in the tropics, and ecological, and therefore evolutionary, constraints on diversity at higher latitudes. For the Bergmann effect, the results support energetic hypotheses that the very largest-bodied and the small-bodied taxa cannot survive the long periods of limited resources at higher latitudes.

Energetics and the evolution of carnivorous plants—Darwin's ‘most wonderful plants in the world’

Carnivory has evolved independently at least six times in five angiosperm orders. In spite of these independent origins, there is a remarkable morphological convergence of carnivorous plant traps and physiological convergence of mechanisms for digesting and assimilating prey. These convergent traits have made carnivorous plants model systems for addressing questions in plant molecular genetics, physiology, and evolutionary ecology. New data show that carnivorous plant genera with morphologically complex traps have higher relative rates of gene substitutions than do those with simple sticky traps. This observation suggests two alternative mechanisms for the evolution and diversification of carnivorous plant lineages. The 'energetics hypothesis' posits rapid morphological evolution resulting from a few changes in regulatory genes responsible for meeting the high energetic demands of active traps. The 'predictable prey capture hypothesis' further posits that complex traps yield more predictable and frequent prey captures. To evaluate these hypotheses, available data on the tempo and mode of carnivorous plant evolution were reviewed; patterns of prey capture by carnivorous plants were analysed; and the energetic costs and benefits of botanical carnivory were re-evaluated. Collectively, the data are more supportive of the energetics hypothesis than the predictable prey capture hypothesis. The energetics hypothesis is consistent with a phenomenological cost-benefit model for the evolution of botanical carnivory, and also accounts for data suggesting that carnivorous plants have leaf construction costs and scaling relationships among leaf traits that are substantially different from those of non-carnivorous plants.

Energetics and space use: intraspecific and interspecific comparisons of movements and home ranges of two Colubrid snakes

1. Energy requirements explain substantial variation in movement and home range size among birds and mammals. This study assesses whether the same is true of snakes by comparing ratsnakes (Elaphe obsoleta) and racers (Coluber constrictor), ecologically similar species whose energy requirements appear to differ substantially (racers > ratsnakes). 2. Over 4 years 22 Elaphe and 16 Coluber were radio-tracked at the same site in Illinois to examine how movement and home ranges varied by sex and season. 3. Coluber moved more often and further per move than Elaphe, resulting in their estimated mean day range being almost four times larger than that of Elaphe (88.0 m day(-1) vs. 23.1 m day(-1)). 4. Both male and female Elaphe moved more frequently early in the season consistent with mate-searching, but mean distances moved did not differ seasonally or by sex. Both sexes of Coluber moved more later in the season and overall males moved further than females. 5. Interspecifically, patterns were consistent with the energetics hypothesis--Coluber had mean home ranges approximately four times larger than those of Elaphe. 6. Intraspecifically, increased movement did not always produce larger home ranges. Male Elaphe had larger home ranges than females despite not moving further, whereas male Coluber had comparable home ranges to females despite moving further. Also, Elaphe home ranges in Illinois were substantially smaller than has been documented in Ontario, despite Ontario Elaphe moving less. 7. Our results generally support the energetics hypothesis, but indicate that knowledge of ecology and energetics increases our understanding of area requirements beyond simple allometric predictions.

Parasitism, life history traits and immune defence in cyprinid fish from Central Europe

BackgroundThe main prediction of life-history theory is that optimal energy allocated among the traits is related to the growth, maintenance and survival. It is hypothesized that the optimal resource allocated to immune function, which generates resistance towards parasites and reduce the fitness losses caused by parasitism, is depending on other requirements for energetic resource and the benefits associated with them. The aims of this study are to investigate in a comparative way (1) how parasitism is related to fish life history traits (fecundity, longevity, mortality), (2) whether there is a trade-off between reproduction and immune investments in fish females (i.e. energetic hypothesis) and in males (i.e. immunohandicap hypothesis), (3) whether parasitism influences host immunity (spleen size) and reproduction (gonad size) in females and males.ResultsData on metazoan parasites of 23 cyprinid fish species from Central Europe were used for the analyses as well as new data collected from a field study. Ectoparasite species richness was negatively correlated with the fish mortality estimated by the k-value and positively correlated with fish body size, suggesting that parasite diversity increases with fish longevity. A negative relationship between spleen size and gonad size, controlling for fish body size, was found in females but not in males. Moreover, parasite abundance was positively correlated with fish spleen size and negatively with fish gonad size in females.ConclusionThe comparative analyses using cyprinid fish species demonstrated that natural mortality could be considered as a factor contributing to the variability of parasite species richness and moreover, parasite species benefit from long-lived fish. The results obtained from the analyses investigating the potential trade-off between reproduction and immunity could be interpreted as an energetic trade-off between female reproduction and immune function. The lack of negative relationship between gonad size and spleen size in males did not support our prediction based on the immunohandicap hypothesis.

MASS LOSS BY BREEDING FEMALE SONGBIRDS: FOOD SUPPLEMENTATION SUPPORTS ENERGETIC STRESS HYPOTHESIS IN BLACK-THROATED BLUE WARBLERS

Abstract Reproduction is physiologically stressful for many animals. Female birds often lose body mass after their young hatch, which has been commonly attributed either to an energetic deficit incurred while breeding (the energetic stress hypothesis) or to an adaptive reduction in wing-loading to save energy during flight (the wing-loading hypothesis). We tested these two hypotheses for an open-cup nesting passerine, the Black-throated Blue Warbler (Dendroica caerulescens), using a food-supplementation experiment. We found that females provided with extra food lost less body mass between the incubation and nestling periods than did control females and that mass loss varied seasonally, with females in both treatments that nested earlier in the season losing more mass. We conclude that greater mass loss in control females supports the energetic stress hypothesis in Black-throated Blue Warblers.

What explains the trot–gallop transition in small mammals?

The transition from trot to gallop in quadruped mammals has been widely hypothesized to be a strategy to minimize the energetic costs of running. This view, however, has been challenged by some experimental evidence suggesting instead that this transition might be triggered by mechanical cues, and would occur when musculoskeletal stresses reach a certain critical value. All previous experiments to test those hypotheses have used relatively large species and their results, therefore, may not be applicable to small mammals. In this study we evaluated the effect of carrying loads on the locomotor energetics and gait transitions of the rodent Octodon degus running on a treadmill. Metabolic rate and cost of transport increased about 30% with a 20% increment in body mass. This increment was higher than expectations based on other mammals, where energy consumption increases in proportion to the added mass, but similar to the response of humans to loads. No abrupt change of energy consumption between gaits was observed and therefore no evidence was found to support the energetic hypothesis. The trot-gallop transition speed did not vary when subjects were experimentally loaded, suggesting that the forces applied to the musculoskeletal system do not trigger gait transition.