Life-history evolution under a production constraint

The recently formulated metabolic theory of ecology has profound implications for the evolution of life histories. Metabolic rate constrains the scaling of production with body mass, so that larger organisms have lower rates of production on a mass-specific basis than smaller ones. Here, we explore the implications of this constraint for life-history evolution. We show that for a range of very simple life histories, Darwinian fitness is equal to birth rate minus death rate. So, natural selection maximizes birth and production rates and minimizes death rates. This implies that decreased body size will generally be favored because it increases production, so long as mortality is unaffected. Alternatively, increased body size will be favored only if it decreases mortality or enhances reproductive success sufficiently to override the preexisting production constraint. Adaptations that may favor evolution of larger size include niche shifts that decrease mortality by escaping predation or that increase fecundity by exploiting new abundant food sources. These principles can be generalized to better understand the intimate relationship between the genetic currency of evolution and the metabolic currency of ecology.

The use of image analysis to estimate population growth rate in Daphnia magna

1. Population growth rate (PGR) is central to the theory of population ecology and is crucial for projecting population trends in conservation biology, pest management and wildlife harvesting. Furthermore, PGR is increasingly used to assess the effects of stressors. Image analysis that can automatically count and measure photographed individuals offers a potential methodology for estimating PGR. 2. This study evaluated two ways in which the PGR of Daphnia magna, exposed to different stressors, can be estimated using an image analysis system. The first method estimated PGR as the ratio of counts of individuals obtained at two different times, while the second method estimated PGR as the ratio of population sizes at two different times, where size is measured by the sum of the individuals' surface areas, i.e. total population surface area. This method is attractive if surface area is correlated with reproductive value (RV), as it is for D. magna, because of the theoretical result that PGR is the rate at which the population RV increases. 3. The image analysis system proved reliable and reproducible in counting populations of up to 440 individuals in 5 L of water. Image counts correlated well with manual counts but with a systematic underestimate of about 30%. This does not affect accuracy when estimating PGR as the ratio of two counts. Area estimates of PGR correlated well with count estimates, but were systematically higher, possibly reflecting their greater accuracy in the study situation. 4. Analysis of relevant scenarios suggested the correlation between RV and body size will generally be good for organisms in which fecundity correlates with body size. In these circumstances, area estimation of PGR is theoretically better than count estimation. 5. Synthesis and applications. There are both theoretical and practical advantages to area estimation of population growth rate when individuals' reproductive values are consistently well correlated with their surface areas. Because stressors may affect both the number and quality of individuals, area estimation of population growth rate should improve the accuracy of predicting stress impacts at the population level.

The allometry of ornaments and weapons

Exaggerated male traits that have evolved under sexual selection include ornaments to attract mates and weapons to deter rivals. Data from studies of many such traits in diverse kinds of organisms show that they almost universally exhibit positive allometries. Both ornaments and weapons increase disproportionately with overall body size, resulting in scaling exponents within species that are consistently > 1.0 and usually in the range 1.5-2.5. We show how scaling exponents reflect the relative fitness advantages of ornaments vs. somatic growth by using a simple mathematical model of resource allocation during ontogeny. Because the scaling exponents are similar for the different taxonomic groups, it follows that the fitness advantages of investing in ornaments also are similar. The model also shows how selection for ornaments influences body size at first reproduction and explains why interspecific allometries have consistently lower exponents than intraspecific ones.

Analytical and compositional aspects of isoflavones in food and their biological effects

This paper provides an overview of analytical techniques used to determine isoflavones (IFs) in foods and biological fluids with main emphasis on sample preparation methods. Factors influencing the content of IFs in food including processing and natural variability are summarized and an insight into IF databases is given. Comparisons of dietary intake of IFs in Asian and Western populations, in special subgroups like vegetarians, vegans, and infants are made and our knowledge on their absorption, distribution, metabolism, and excretion by the human body is presented. The influences of the gut microflora, age, gender, background diet, food matrix, and the chemical nature of the IFs on the metabolism of IFs are described. Potential mechanisms by which IFs may exert their actions are reviewed, and genetic polymorphism as determinants of biological response to soy IFs is discussed. The effects of IFs on a range of health outcomes including atherosclerosis, breast, intestinal, and prostate cancers, menopausal symptoms, bone health, and cognition are reviewed on the basis of the available in vitro, in vivo animal and human data.

ICT implants: the invasive future of identity?

Is the human body a suitable place for a microchip? Such discussion is no longer hypothetical - in fact in reality it has not been so for some years. Restorative devices such as pacemakers and cochlear implants have become well established, yet these sophisticated devices form notably intimate links between technology and the body. More recent developments in engineering technologies have meant that the integration of silicon with biology is now reaching new levels - with devices which interact directly with the brain. As medical technologies continue to advance, their potential benefits for human enhancement will become increasingly attractive, and so we need to seriously consider where this may take us. In this paper, an attempt is made to demonstrate that, in the medical context, the foundations of more advanced implantable enhancement technologies are already notably progressed, and that they are becoming more science fact than is widely considered. A number of wider moral, ethical and legal issues stem from enhancement applications and it is difficult to foresee the social consequences, the fundamental changes on our very conception of self and the impact on our identity of adoption long term. As a result, it is necessary to acknowledge the possibilities and is timely to have debate to address the wider implications these possibilities may bring.

Temperature and the development rates of thrips: evidence for a constraint on local adaptation?

Typically, the relationship between insect development and temperature is described by two characteristics: the minimum temperature needed for development to occur (T-min) and the number of day degrees required (DDR) for the completion of development. We investigated these characteristics in three English populations of Thrips major and T tabaci [Cawood, Yorkshire (N53degrees49', W1degrees7'); Boxworth, Cambridgeshire (N52degrees15', W0degrees1'); Silwood Park, Berkshire (N51degrees24', W0degrees38')], and two populations of Frankliniella occidentalis (Cawood; Silwood Park). While there were no significant differences among populations in either T-min (mean for T major = 7.0degreesC; T tabaci = 5.9degreesC; F. occidentalis = 6.7degreesC) or DDR (mean for T major = 229.9; T tabaci = 260.8; F occidentalis = 233.4), there were significant differences in the relationship between temperature and body size, suggesting the presence of geographic variation in this trait. Using published data, in addition to those newly collected, we found a negative relationship between T-min. and DDR for F occidentalis and T tabaci, supporting the hypothesis that a trade-off between T-min and DDR may constrain adaptation to local climatic conditions.

Phylogeny and metabolic scaling in mammals

The scaling of metabolic rates to body size is widely considered to be of great biological and ecological importance, and much attention has been devoted to determining its theoretical and empirical value. Most debate centers on whether the underlying power law describing metabolic rates is 2/3 (as predicted by scaling of surface area/volume relationships) or 3/4 ("Kleiber's law"). Although recent evidence suggests that empirically derived exponents vary among clades with radically different metabolic strategies, such as ectotherms and endotherms, models, such as the metabolic theory of ecology, depend on the assumption that there is at least a predominant, if not universal, metabolic scaling exponent. Most analyses claimed to support the predictions of general models, however, failed to control for phylogeny. We used phylogenetic generalized least-squares models to estimate allometric slopes for both basal metabolic rate (BMR) and field metabolic rate (FMR) in mammals. Metabolic rate scaling conformed to no single theoretical prediction, but varied significantly among phylogenetic lineages. In some lineages we found a 3/4 exponent, in others a 2/3 exponent, and in yet others exponents differed significantly from both theoretical values. Analysis of the phylogenetic signal in the data indicated that the assumptions of neither species-level analysis nor independent contrasts were met. Analyses that assumed no phylogenetic signal in the data (species-level analysis) or a strong phylogenetic signal (independent contrasts), therefore, returned estimates of allometric slopes that were erroneous in 30% and 50% of cases, respectively. Hence, quantitative estimation of the phylogenetic signal is essential for determining scaling exponents. The lack of evidence for a predominant scaling exponent in these analyses suggests that general models of metabolic scaling, and macro-ecological theories that depend on them, have little explanatory power.

Exposure to environmental oestrogenic chemicals and breast cancer.

The human population is now exposed on a daily basis to a multitude of environmental pollutant chemicals that would not have been present a century ago, and many of these chemicals have been detected in the human breast. The fatty nature of human breast tissue makes it a particular target for lipophilic as well as hydrophilic pollutant chemicals, which may enter the human body through oral, respiratory, or dermal routes. These chemicals possess a range of endocrine-disrupting properties and genotoxic activity, but from a breast cancer perspective the greatest concern has centered around their ability to mimic or interfere with the action of estrogen. The breast is an endocrine target organ and exposure to estrogen is a known risk factor for breast cancer.

Advances in mucoadhesion and mucoadhesive polymers

Mucoadhesion is the ability of materials to adhere to mucosal membranes in the human body and provide a temporary retention. This property has been widely used to develop polymeric dosage forms for buccal, oral, nasal, ocular and vaginal drug delivery. Excellent mucoadhesive properties are typical for hydrophilic polymers possessing charged groups and/or non-ionic functional groups capable of forming hydrogen bonds with mucosal surfaces. This feature article considers recent advances in the study of mucoadhesion and mucoadhesive polymers. It provides an overview on the structure of mucosal membranes, properties of mucus gels and the nature of mucoadhesion. It describes the most common methods to evaluate mucoadhesive properties of various dosage forms and discusses the main classes of mucoadhesives.

The ecology of lizard reproductive output

Aim  We provide a new quantitative analysis of lizard reproductive ecology. Comparative studies of lizard reproduction to date have usually considered life-history components separately. Instead, we examine the rate of production (productivity hereafter) calculated as the total mass of offspring produced in a year. We test whether productivity is influenced by proxies of adult mortality rates such as insularity and fossorial habits, by measures of temperature such as environmental and body temperatures, mode of reproduction and activity times, and by environmental productivity and diet. We further examine whether low productivity is linked to high extinction risk. Location  World-wide. Methods  We assembled a database containing 551 lizard species, their phylogenetic relationships and multiple life history and ecological variables from the literature. We use phylogenetically informed statistical models to estimate the factors related to lizard productivity. Results  Some, but not all, predictions of metabolic and life-history theories are supported. When analysed separately, clutch size, relative clutch mass and brood frequency are poorly correlated with body mass, but their product – productivity – is well correlated with mass. The allometry of productivity scales similarly to metabolic rate, suggesting that a constant fraction of assimilated energy is allocated to production irrespective of body size. Island species were less productive than continental species. Mass-specific productivity was positively correlated with environmental temperature, but not with body temperature. Viviparous lizards were less productive than egg-laying species. Diet and primary productivity were not associated with productivity in any model. Other effects, including lower productivity of fossorial, nocturnal and active foraging species were confounded with phylogeny. Productivity was not lower in species at risk of extinction. Main conclusions  Our analyses show the value of focusing on the rate of annual biomass production (productivity), and generally supported associations between productivity and environmental temperature, factors that affect mortality and the number of broods a lizard can produce in a year, but not with measures of body temperature, environmental productivity or diet.

Shifts in metabolic scaling, production, and efficiency across major evolutionary transitions of life

The diversification of life involved enormous increases in size and complexity. The evolutionary transitions from prokaryotes to unicellular eukaryotes to metazoans were accompanied by major innovations inmetabolicdesign.Hereweshowthat thescalingsofmetabolic rate, population growth rate, and production efficiency with body size have changed across the evolutionary transitions.Metabolic rate scales with body mass superlinearly in prokaryotes, linearly in protists, and sublinearly inmetazoans, so Kleiber’s 3/4 power scaling law does not apply universally across organisms. The scaling ofmaximum population growth rate shifts from positive in prokaryotes to negative in protists and metazoans, and the efficiency of production declines across these groups.Major changes inmetabolic processes duringtheearlyevolutionof life overcameexistingconstraints, exploited new opportunities, and imposed new constraints. The 3.5 billion year history of life on earth was characterized by

Experiments into biology-technology interaction

In this paper a look is taken at how the use of implant and electrode technology can be employed to create biological brains for robots, to enable human enhancement and to diminish the effects of certain neural illnesses. In all cases the end result is to increase the range of abilities of the recipients. An indication is given of a number of areas in which such technology has already had a profound effect, a key element being the need for a clear interface linking a biological brain directly with computer technology. The emphasis is placed on practical scientific studies that have been and are being undertaken and reported on. The area of focus is the use of electrode technology, where either a connection is made directly with the cerebral cortex and/or nervous system or where implants into the human body are involved. The paper also considers robots that have biological brains in which human neurons can be employed as the sole thinking machine for a real world robot body.

Are patterns of growth adaptive?

Models which define fitness in terms of per capita rate of increase of phenotypes are used to analyse patterns of individual growth. It is shown that sigmoid growth curves are an optimal strategy (i.e. maximize fitness) if (Assumption 1a) mortality decreases with body size; (2a) mortality is a convex function of specific growth rate, viewed from above; (3) there is a constraint on growth rate, which is attained in the first phase of growth. If the constraint is not attained then size should increase at a progressively reducing rate. These predictions are biologically plausible. Catch-up growth, for retarded individuals, is generally not an optimal strategy though in special cases (e.g. seasonal breeding) it might be. Growth may be advantageous after first breeding if birth rate is a convex function of G (the fraction of production devoted to growth) viewed from above (Assumption 5a), or if mortality rate is a convex function of G, viewed from above (Assumption 6c). If assumptions 5a and 6c are both false, growth should cease at the age of first reproduction. These predictions could be used to evaluate the incidence of indeterminate versus determinate growth in the animal kingdom though the data currently available do not allow quantitative tests. In animals with invariant adult size a method is given which allows one to calculate whether an increase in body size is favoured given that fecundity and developmental time are thereby increased.

Rensch's rule in large herbivorous mammals derived from metabolic scaling

Rensch’s rule, which states that the magnitude of sexual size dimorphism tends to increase with increasing body size, has evolved independently in three lineages of large herbivorous mammals: bovids (antelopes), cervids (deer), and macropodids (kangaroos). This pattern can be explained by a model that combines allometry,life-history theory, and energetics. The key features are thatfemale group size increases with increasing body size and that males have evolved under sexual selection to grow large enough to control these groups of females. The model predicts relationships among body size and female group size, male and female age at first breeding,death and growth rates, and energy allocation of males to produce body mass and weapons. Model predictions are well supported by data for these megaherbivores. The model suggests hypotheses for why some other sexually dimorphic taxa, such as primates and pinnipeds(seals and sea lions), do or do not conform to Rensh’s rule.

Energetics, lifestyle, and reproduction in birds

Theoretical and empirical studies of life history aim to account for resource allocation to the different components of fitness: survival, growth, and reproduction. The pioneering evolutionary ecologist David Lack [(1968) Ecological Adaptations for Breeding in Birds (Methuen and Co.,London)] suggested that reproductive output in birds reflects adaptation to environmental factors such as availability of food and risk of predation, but subsequent studies have not always supported Lack’s interpretation. Here using a dataset for 980 bird species (Dataset S1), a phylogeny, and an explicit measure of reproductive productivity, we test predictions for how mass-specific productivity varies with body size, phylogeny,and lifestyle traits. We find that productivity varies negatively with body size and energetic demands of parental care and positively with extrinsic mortality. Specifically: (i) altricial species are 50% less productive than precocial species; (ii) species with female-only care of offspring are about 20% less productive than species with other methods of parental care; (iii) nonmigrants are 14% less productive than migrants; (iv) frugivores and nectarivores are about 20% less productive than those eating other foods; and (v) pelagic foragers are 40% less productive than those feeding in other habitats. A strong signal of phylogeny suggests that syndromes of similar life-history traits tend to be conservative within clades but also to have evolved independently in different clades. Our results generally support both Lack’s pioneering studies and subsequent research on avian life history.