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Group selection, altruism, reinforcement, and throwing in human evolution.

Evolution of altruism by group selection involves sacrifice of some individuals, not to the "group as a whole," but to other individuals in the group. Deme-group selection may establish strictly altruistic genes in a population, but only under limited conditions, and perhaps never among vertebrates, among which apparently altruistic behaviors may always potentially benefit the altruists. Responsive-group selection is a more effective mode of evolution of altruism, conspicuous in man. Evolutionary reinforcement increases the force of selection of advantageous behaviors, including altruistic ones, by making them pleasant or rewarding. It is probably involved also in ecological habitat selection, and may be the source of many human emotions, including esthetic ones. Throwing (of stones and weapons) exemplifies both the possible importance of a difficult-to-measure evolutionary factor and the role of reinforcement; in human evolution throwing may have been decisive in food-getting and fighting, in shifting emphasis from brute force to skill, and in inducing evolution of a brain able to handle three-body geometric problems precisely and thus preadapted for more complex functions.

Anthropology, Physical↗

Predatory prokaryotes: predation and primary consumption evolved in bacteria.

Two kinds of predatory bacteria have been observed and characterized by light and electron microscopy in samples from freshwater sulfurous lakes in northeastern Spain. The first bacterium, named Vampirococcus, is Gram-negative and ovoidal (0.6 micrometer wide). An anaerobic epibiont, it adheres to the surface of phototrophic bacteria (Chromatium spp.) by specific attachment structures and, as it grows and divides by fission, destroys its prey. An important in situ predatory role can be inferred for Vampirococcus from direct counts in natural samples. The second bacterium, named Daptobacter, is a Gram-negative, facultatively anaerobic straight rod (0.5 x 1.5 micrometers) with a single polar flagellum, which collides, penetrates, and grows inside the cytoplasm of its prey (several genera of Chromatiaceae). Considering also the well-known case of Bdellovibrio, a Gram-negative, aerobic curved rod that penetrates and divides in the periplasmic space of many chemotrophic Gram-negative bacteria, there are three types of predatory prokaryotes presently known (epibiotic, cytoplasmic, and periplasmic). Thus, we conclude that antagonistic relationships such as primary consumption, predation, and scavenging had already evolved in microbial ecosystems prior to the appearance of eukaryotes. Furthermore, because they represent methods by which prokaryotes can penetrate other prokaryotes in the absence of phagocytosis, these associations can be considered preadaptation for the origin of intracellular organelles.

Bdellovibrio↗

Latent capacities for gametogenic cycling in the semelparous invertebrate Nereis.

Most nereid polychaetes are strictly semelparous, a single episode of reproduction being invariably followed by death. Endocrine manipulation in Nereis diversicolor by the regular implantation of cerebral ganglia from immature donors unveils characteristics associated with a capacity to engage in repeated gametogenic cycling. Such manipulation permits full maturation of the gametes but blocks spawning. Gamete resorption then leads on to another bout of gametogenesis and a new cohort of gametes is formed. The neurosecretory system adopts a cyclical pattern of activity, which parallels that of gametogenesis. Repair and maintenance of the soma continue throughout sexual maturation, as shown by the persistence of feeding and the capacity for regenerative segment proliferation. In consequence, life is extended apparently indefinitely. These latent capacities are reminiscent of features of iteroparous life histories, characterized by repeated breeding, and are postulated to be vestiges of an iteroparous ancestry. They also constitute a preadaptation for iteroparity and reveal how readily a reversal to this condition could occur. The study suggests that reproductive strategies may be unexpectedly labile in even their most fundamental aspects.

Aging↗

Heat shock protein synthesis and thermotolerance in Cataglyphis, an ant from the Sahara desert.

The ant Cataglyphis lives in the Sahara desert and is one of the most thermotolerant land animals known. It forages at body temperatures above 50 degrees C, and the critical thermal maxima are at 53.6 +/- 0.8 degrees C for Cataglyphis bombycina and 55.1 +/- 1.1 degrees C for Cataglyphis bicolor. The synthesis and accumulation of heat shock proteins (HSPs) were analyzed in Cataglyphis and compared to Formica, an ant living in more moderate climates, and to two Drosophila species. In Cataglyphis, protein synthesis continues at temperatures up to 45 degrees C as compared to 39 degrees C for Formica and Drosophila. The two Drosophila species, Drosophila melanogaster and Drosophila ambigua, differ with respect to their maximal induction of HSP synthesis and accumulation by 3-4 degrees C. In contrast, the two ant species accumulate HSPs prior to their exposure to heat, and in Cataglyphis the temperature of maximal HSP induction by de novo protein synthesis is only 2 degrees C higher than in Formica. These findings are interpreted as preadaption of the ants prior to exposure to high temperatures.

Acclimatization↗

Evolutionary relationships of the coelacanth, lungfishes, and tetrapods based on the 28S ribosomal RNA gene.

The origin of land vertebrates was one of the major transitions in the history of vertebrates. Yet, despite many studies that are based on either morphology or molecules, the phylogenetic relationships among tetrapods and the other two living groups of lobe-finned fishes, the coelacanth and the lungfishes, are still unresolved and debated. Knowledge of the relationships among these lineages, which originated back in the Devonian, has profound implications for the reconstruction of the evolutionary scenario of the conquest of land. We collected the largest molecular data set on this issue so far, about 3,500 base pairs from seven species of the large 28S nuclear ribosomal gene. All phylogenetic analyses (maximum parsimony, neighbor-joining, and maximum likelihood) point toward the hypothesis that lungfishes and coelacanths form a monophyletic group and are equally closely related to land vertebrates. This evolutionary hypothesis complicates the identification of morphological or physiological preadaptations that might have permitted the common ancestor of tetrapods to colonize land. This is because the reconstruction of its ancestral conditions would be hindered by the difficulty to separate uniquely derived characters from shared derived characters in the coelacanth/lungfish and tetrapod lineages. This molecular phylogeny aids in the reconstruction of morphological evolutionary steps by providing a framework; however, only paleontological evidence can determine the sequence of morphological acquisitions that allowed lobe-finned fishes to colonize land.

Animals↗

Chemical chaperones regulate molecular chaperones in vitro and in cells under combined salt and heat stresses.

Salt and heat stresses, which are often combined in nature, induce complementing defense mechanisms. Organisms adapt to high external salinity by accumulating small organic compounds known as osmolytes, which equilibrate cellular osmotic pressure. Osmolytes can also act as "chemical chaperones" by increasing the stability of native proteins and assisting refolding of unfolded polypeptides. Adaptation to heat stress depends on the expression of heat-shock proteins, many of which are molecular chaperones, that prevent protein aggregation, disassemble protein aggregates, and assist protein refolding. We show here that Escherichia coli cells preadapted to high salinity contain increased levels of glycine betaine that prevent protein aggregation under thermal stress. After heat shock, the aggregated proteins, which escaped protection, were disaggregated in salt-adapted cells as efficiently as in low salt. Here we address the effects of four common osmolytes on chaperone activity in vitro. Systematic dose responses of glycine betaine, glycerol, proline, and trehalose revealed a regulatory effect on the folding activities of individual and combinations of chaperones GroEL, DnaK, and ClpB. With the exception of trehalose, low physiological concentrations of proline, glycerol, and especially glycine betaine activated the molecular chaperones, likely by assisting local folding in chaperone-bound polypeptides and stabilizing the native end product of the reaction. High osmolyte concentrations, especially trehalose, strongly inhibited DnaK-dependent chaperone networks, such as DnaK+GroEL and DnaK+ClpB, likely because high viscosity affects dynamic interactions between chaperones and folding substrates and stabilizes protein aggregates. Thus, during combined salt and heat stresses, cells can specifically control protein stability and chaperone-mediated disaggregation and refolding by modulating the intracellular levels of different osmolytes.

Adaptation, Biological↗

Gender Differences in Intercepting a Moving Target by Using a Throw or Button Press.

Humans are a throwing species just as much as a talking species (Calvin, 1982), but preadaptations for throwing remain largely unexplored. The ability of undergraduate students (N = 165) to intercept a stimulus with a thrown object was determined for 10 stimulus speeds, ranging from 2 to 5 m/s. The inclusion of a condition in which the moving stimulus was intercepted via button press allowed the role of the perceptual aspect of velocity estimation to be evaluated separately from the act of throwing. Gender differences were observed in both the button and the throwing intercept tasks: The men had smaller timing errors and higher accuracy than the women. Contrary to expectations, the male and female patterns of intercept errors as a function of stimulus speed were more similar in the throwing task than in the button intercept task.

gender differences↗

Chronic dietary restriction influences tumor metastasis in the rat: parametric considerations.

Chronic dietary restriction is a well-documented means of inhibiting tumor growth. This study examines the effects of chronic dietary restriction on tumor metastasis in the rat. We investigate the effect of 1) the degree of food restriction, 2) the effect of preexposure to food restriction, and 3) the duration of food restriction after tumor inoculation on tumor metastasis. We also compare two methods of dietary restriction: 1) the time that food is available and 2) the amount of food available. Our findings demonstrate that rats restricted to 50% of ad libitum diet for one week before inoculation with MADB106 tumor cells and for three weeks after inoculation exhibit a significant (p < 0.001) reduction in lung colonization compared with animals fed ad libitum. Animals restricted to access to food for 4 hrs/day (60% of ad libitum) for the same period of time exhibit significantly (p < 0.005) greater lung tumor colonization than animals fed ad libitum. Preadaptation to the feeding regimen for one week before tumor inoculation proved to be critical in inhibiting tumor metastasis. The tumor-inhibitory effect was not significantly influenced by the duration of restriction after inoculation or by the manner in which food restriction was implemented. Finally, we demonstrate that inhibition of tumor colonization may be mediated by enhanced natural killer cell activity in the early postinoculation period.

Adenocarcinoma↗

Toxicokinetics and effects of PCBs in Arctic fish: a review of studies on Arctic charr.

In a series of environmentally realistic laboratory experiments, toxicokinetics and effects of polychlorinated biphenyls (PCBs) were studied in the Arctic charr (Salvelinus alpinus). Winter fasting and emaciation, which are common among Arctic charr living in high latitudes, resulted in a redistribution of the lipophilic PCBs from lipid-storing tissue such as the muscle, to vital organs that must be considered sensitive toward PCB (liver and brain). This redistribution was accompanied by a significant potentiation of the hepatic cytochrome P-450 (CYP) 1A biomarker response, from low activities in October (within those measured in uncontaminated charr) to a high, probably maximum, induction in May. Performance studies demonstrated a clear effect of environmentally realistic PCB levels on endocrine mechanisms, immune function, and seawater preadaptation (smoltification) in charr that had been feed deprived for several months after contamination with Aroclor 1254, whereas a high PCB dose exerted only minor, if any, effects in charr that had been fed after contamination. These results demonstrate that emaciation results in decreased dose-response relationships in fish, and indicate that arctic animals undergoing seasonal cycles of "fattening" and emaciation may be extra sensitive toward persistent, lipophilic organochlorines. Pilot studies on Arctic charr from Bjørnøya Island revealed marked CYP1A biomarker responses and an upregulation of genes involved in cellular homeostatic mechanisms in charr from Lake Ellasjøen (high PCB levels).

Adaptation, Physiological↗

The broad spectrum of responses to oxidants in proliferating cells: a new paradigm for oxidative stress.

Proliferating mammalian cells exhibit a broad spectrum of responses to oxidative stress, depending on the stress level encountered. Very low levels of hydrogen peroxide, e.g., 3 to 15 microM, or 0.1 to 0.5 micromol/10(7) cells, cause a significant mitogenic response, 25% to 45 % growth stimulation. Greater concentrations of H2O2, 120 to 150 microM, or 2 to 5 micromol/10(7) cells, cause a temporary growth arrest that appears to protect cells from excess energy use and DNA damage. After 4-6 h of temporary growth arrest, many cells will exhibit up to a 40-fold transient adaptive response in which genes for oxidant protection and damage repair are preferentially expressed. After 18 h of H2O2 adaptation (including the 4-6 h of temporary growth arrest) cells exhibit maximal protection against oxidative stress. The H2O2 originally added is metabolized within 30-40 min, and if no more is added the cells will gradually de-adapt, so that by 36 h after the initial H2O2 stimulus they have returned to their original level of H2O2 sensitivity. At H2O2 concentrations of 250 to 400 microM, or 9 to 14 micromol/10(7) cells, mammalian fibroblasts are not able to adapt but instead enter a permanently growth-arrested state in which they appear to perform most normal cell functions but never divide again. This state of permanent growth arrest has often been confused with cell death in toxicity studies relying solely on cell proliferation assays as measures of viability. If the oxidative stress level is further increased to 0.5 to 1.0 mM H2O2, or 15 to 30 micromol/10(7) cells, apoptosis results. This oxidative stress-induced apoptosis involves nuclear condensation, loss of mitochondrial transmembrane potential, degradation/down-regulation of mitochondrial mRNAs and rRNAs, and degradation/laddering of both nuclear and mitochondrial DNA. At very high H2O2 concentrations of 5.0 to 10.0 mM, or 150 to 300 micromol/10(7) cells and above, cell membranes disintegrate, proteins and nucleic acids denature, and necrosis swiftly follows. Cultured cells grown in 20% oxygen are essentially preadapted or preselected to survive under conditions of oxidative stress. If cells are instead grown in 3% oxygen, much closer to physiological cellular levels, they are more sensitive to an oxidative challenge but exhibit far less accumulated oxidant damage. This broad spectrum of cellular responses to oxidant stress, depending on the amount of oxidant applied and the concentration of oxygen in the cell culture system, provides for a new paradigm of cellular oxidative stress responses.

Animals↗

The nature of the in vivo sodium and chloride uptake mechanisms through the epithelium against sodium and of bicarbonate against chloride.

The Chilean frog, Calyptocephallela gayi, placed in dilute NaCl solutions may pump Na(+) and Cl(-) at very different rates depending on the kind of bath solutions in which it was preadapted. Furthermore, Na(+) and Cl(-) may be absorbed from solutions in which the accompanying coion, such as sulfate and choline, respectively, is impermeant. In all these cases it is obligatory to postulate the existence of two ionic exchange mechanisms, Cl(-) and Na(+), being exchanged against endogenous anions and cations, respectively. It has been determined that Na(+) is exchanged against endogenous H(+) and that Cl(-) is exchanged against HCO(3) (-). In animals pumping Na(+) and Cl(-) from dilute NaCl solutions Na(+) or Cl(-) uptake may be selectively inhibited, while the flux of the accompanying ion remains unchanged. This is considered to be an additional proof that both Na(+) and Cl(-) fluxes are always independent. The role of the ionic exchange mechanisms in the direct regulation of the Na(+) and Cl(-) levels in the internal medium is discussed as well as their relationship in the regulation of the acid-base equilibrium; other physioecological considerations have been treated.

Acid-Base Equilibrium↗

Resource allocation in offspring provisioning: an evaluation of the conditions favoring the evolution of matrotrophy.

We used analytic and simulation models to determine the ecological conditions favoring evolution of a matrotrophic fish from a lecithotrophic ancestor given a complex set of trade-offs. Matrotrophy is the nourishment of viviparous embryos by resources provided between fertilization and parturition, while lecithotrophy describes embryo nourishment provided before fertilization. In fishes and reptiles, embryo nourishment encompasses a continuum from solely lecithotrophic to primarily matrotrophic. Matrotrophy has evolved independently from lecithotrophic ancestors many times in many groups. We assumed matrotrophy increased the number of offspring a viviparous female could gestate and evaluated conditions of food availability favoring lecithotrophy or matrotrophy. The matrotrophic strategy was superior when food resources exceeded demand during gestation but at a risk of overproduction and reproductive failure if food intake was limited. Matrotrophic females were leaner during gestation than lecithotrophic females, yielding shorter life spans. Our models suggest that matrotrophic embryo nourishment evolved in environments with high food availability, consistently exceeding energy requirements for maintaining relatively large broods. Embryo abortion with some resorption of invested energy is a necessary preadaptation to the evolution of matrotrophy. Future work should explore trade-offs of age-specific mortality and reproductive output for females maintaining different levels of fat storage during gestation.

Animals↗

Maternal effects on offspring locomotion: influence of density and corticosterone elevation in the lizard Lacerta vivipara.

Offspring phenotype can be affected by maternal history before and during gestation. Offspring sensitivity to maternal conditions is believed to have evolved to favor preadaptation of offspring to environmental factors they are likely to encounter. Because the locomotor capacity of an individual is likely to have important fitness consequences, we examined the role of long-term and short-term prenatal conditions on offspring's locomotor performance in the lizard Lacerta vivipara. To examine long-term prenatal effects, we manipulated the density of two populations, leaving two additional populations as unmanipulated. We then collected pregnant females within these four populations (Cévennes, Massif Central, France) and kept them in the laboratory until parturition. To examine short-term prenatal effects, we manipulated the corticosterone level of half the females within each population. We took two different measurements of offspring locomotion: sprint speed and endurance. As already documented, sprint speed was positively correlated with offspring body size. Although population density significantly affected female fecundity, neither the density manipulation nor the population of origin influenced offspring phenotype. Corticosterone administered during gestation decreased juvenile sprint speed but did not affect juvenile endurance. Furthermore, we observed that the motivation to run was influenced by maternal hormonal treatment. Juveniles born from corticosterone-treated mothers needed more stimuli than those born from control mothers. We conclude, therefore, that the action of corticosterone on sprint speed could be more behavioral than physiological. Offspring phenotype as measured by endurance and sprint speed appeared partly under maternal control.

Animals↗

The evolutionary origin of feathers.

Previous theories relating the origin of feathers to flight or to heat conservation are considered to be inadequate. There is need for a model of feather evolution that gives attention to the function and adaptive advantage of intermediate structures. The present model attempts to reveal and to deal with, the spectrum of complex questions that must be considered. In several genera of modern lizards, scales are elongated in warm climates. It is argued that these scales act as small shields to solar radiation. Experiments are reported that tend to confirm this. Using lizards as a conceptual model, it is argued that feathers likewise arose as adaptations to intense solar radiation. Elongated scales are assumed to have subdivided into finely branched structures that produced a heat-shield, flexible as well as long and broad. Associated muscles had the function of allowing the organism fine control over rates of heat gain and loss: the specialized scales or early feathers could be moved to allow basking in cool weather or protection in hot weather. Subdivision of the scales also allowed a close fit between the elements of the insulative integument. There would have been mechanical and thermal advantages to having branches that interlocked into a pennaceous structure early in evolution, so the first feathers may have been pennaceous. A versatile insulation of movable, branched scales would have been a preadaptation for endothermy. As birds took to the air they faced cooling problems despite their insulative covering because of high convective heat loss. Short glides may have initially been advantageous in cooling an animal under heat stress, but at some point the problem may have shifted from one of heat exclusion to one of heat retention. Endothermy probably evolved in conjunction with flight. If so, it is an unnecessary assumption to postulate that the climate cooled and made endothermy advantageous. The development of feathers is complex and a model is proposed that gives attention to the fundamental problems of deriving a branched structure with a cylindrical base from an elongated scale.

Animals↗

Termite digestibility and water and energy contents determine the water economy index of numbats (Myrmecobius fasciatus) and other myrmecophages.

Digestibility by captive numbats for termites was determined by feeding trials to be 81%+/-1.2% for Coptotermes sp. and 64%+/-3.3% for Nasutitermes sp. Water, ash, and energy content of both the Coptotermes (0.96+/-0.099 mg(dry mass) individual(-1), 78.0%+/-0.36% water, 5.8%+/-0.31% ash, 23.1+/-0.19 kJ g-1dry total energy) and Nasutitermes (0.91+/-0.046 mg(dry mass) individual(-1), 76.7%+/-3.09% water, 7.5%+/-1.10% ash, 22.7+/-0.36 kJ g-1dry total energy) were similar to values measured previously for other termites and for ants and insects in general. Numbats have a slow passage time for termites (20-30 h), presumably to enhance the digestion of termites. The water economy index (WEI) was 0.2 for captive numbats feeding on Coptotermes and 0.25 for Nasutitermes, whereas the WEI measured for wild, free-living numbats was 0.29, which corresponds to a digestibility of 58%. The WEI of a myrmecophage diet is determined by the energy and water contents and digestibilities of termites and ants, in the absence of drinking. The WEI for numbats, and other termitivorous mammals as well as reptiles, is higher than would be expected for an animal-based diet because of their relatively low digestibility (58%-81%) for termites. A high WEI preadapts myrmecophages to survival in arid environments without having to drink.

Analysis of Variance↗

The evolution and functions of laughter and humor: a synthetic approach.

A number of recent hypotheses have attempted to explain the ultimate evolutionary origins of laughter and humor. However most of these have lacked breadth in their evolutionary frameworks while neglecting the empirical existence of two distinct types of laughter--Duchenne and non-Duchenne--and the implications of this distinction for the evolution of laughter as a signal. Most of these hypotheses have also been proposed in relative isolation of each other and remain disjointed from the relevant empirical literature. Here we attempt to remedy these shortcomings through a synthesis of previous laughter and humor research followed by (i) a reevaluation of this research in light of theory and data from several relevant disciplines, and (ii) the proposal of a synthetic evolutionary framework that takes into account phylogeny and history as well as proximate mechanisms and adaptive significance. We consider laughter to have been a preadaptation that was gradually elaborated and co-opted through both biological and cultural evolution. We hypothesize that Duchenne laughter became fully ritualized in early hominids between 4 and 2 mya as a medium for playful emotional contagion. This mechanism would have coupled the emotions of small hominid groups and promoted resource-building social play during the fleeting periods of safety and satiation that characterized early bipedal life. We further postulate that a generalized class of nonserious social incongruity would have been a reliable indicator of such safe times and thereby came to be a potent distal elicitor of laughter and playful emotion. This class of stimuli had its origins in primate social play and was the foundation for formal human humor. Within this framework, Duchenne laughter and protohumor were well established in the hominid biobehavioral repertoire when more cognitively sophisticated traits evolved in the hominid line between 2 mya and the present. The prior existence of laughter and humor allowed them to be co-opted for numerous novel functions, and it is from this process that non-Duchenne laughter and the "dark side" of laughter emerged. This perspective organizes the diversified forms and functions that characterize laughter and humor today and clarifies when and how laughter and humor evolved during the course of human evolution.

Adaptation, Physiological↗

Production of milk and nutrition of the dependent young of free-ranging koalas (Phascolarctos cinereus).

The production of milk by lactating females, and energy expenditure and foliage intake of their dependent young, were investigated in free-ranging koalas. Koalas had the lowest mass-specific daily milk-energy production at peak lactation so far recorded in a mammal, but the duration of reproduction was 58% longer than the combined marsupial and eutherian average. As a consequence, the total energy input to reproduction in koalas was similar to that in other mammals. We propose that the prolonged lactation and low daily rate of energy transfer to the young by female koalas is an adaptation to the low energy availability from their diet of Eucalyptus foliage. Energy requirements (field metabolic rates) of young koalas were lower than those expected for typical marsupials (only 60% at permanent pouch exit), which may be a necessary preadaptation that allows the low rate of maternal energy transfer. However, the energy requirements of the adult females were no lower than expected for marsupials. This pattern of energy requirements and age resulted in a linear relationship between field metabolic rate and mass for the koalas in this population. Differences in milk production between the years of the study coincided with fluctuations in the availability of preferred young foliage, which suggests that lactational output by koalas may be flexible and affected by diet quality. Despite the interannual differences in milk production, growth of the young was similar in the two years.

Aging↗

Heart rate variability in rats acclimatized to high altitude.

The aims of this study were to relate heart morphology and functions changes to heart rate variability (HRV) components after acclimatization to high altitude and to define whether preadaptation to hypoxia could modulate HRV responses to acute hypoxic stress. Doppler-echocardiographic studies of the left ventricle were performed in female Wistar rats before, during, and after a 10-week exposure to moderate hypobaric hypoxia (CH rats, approximately 4000 m simulated) or normoxia (N rats, approximately 55 m). Right ventricular morphology and function and pulmonary artery pressure were evaluated using heart catheterization. Spectral analysis of HRV was studied after exposure in conscious unrestrained rats in normoxia and during acute hypoxic stress. Necropsy right ventricular hypertrophy and intraventricular and pulmonary artery hypertension were found in CH rats compared with N rats. Echocardiographic left ventricular morphology and functions were similar between the groups after exposures. Compared to the control group, CH rats had similar heart rates and HRV components when measured in normoxia. During acute hypoxic stress, HRV decreased in all rats, but less in CH rats. These results support the hypothesis that long-term mild hypoxia may moderate sympathetic activation induced by acute hypoxia and that right ventricular hypertrophy cannot be the direct cause of such a shift in sympathovagal nerve interaction during acute hypoxic stress.

Acclimatization↗