Peto's paradox and human cancers

Peto''s paradox is the lack of the expected trend in cancer incidence as a function of body size and lifespan across species. The leading hypothesis to explain this pattern is natural selection for differential cancer prevention in larger, longer lived species. We evaluate whether a similar effect exists within species, specifically humans. We begin by reanalysing a recently published dataset to separate the effects of stem cell number and replication rate, and show that each has an independent effect on cancer risk. When considering the lifetime number of stem cell divisions in an extended dataset, and removing cases associated with other diseases or carcinogens, we find that lifetime cancer risk per tissue saturates at approximately 0.3–1.3% for the types considered. We further demonstrate that grouping by anatomical site explains most of the remaining variation. Our results indicate that cancer risk depends not only on the number of stem cell divisions but varies enormously (approx. 10 000 times) depending on anatomical site. We conclude that variation in risk of human cancer types is analogous to the paradoxical lack of variation in cancer incidence among animal species and may likewise be understood as a result of evolution by natural selection.     

The multiple facets of Peto's paradox: a life-history model for the evolution of cancer suppression

Large animals should have higher lifetime probabilities of cancer than small animals because each cell division carries an attendant risk of mutating towards a tumour lineage. However, this is not observed—a (Peto''s) paradox that suggests large and/or long-lived species have evolved effective cancer suppression mechanisms. Using the Euler–Lotka population model, we demonstrate the evolutionary value of cancer suppression as determined by the ‘cost’ (decreased fecundity) of suppression verses the ‘cost’ of cancer (reduced survivorship). Body size per se will not select for sufficient cancer suppression to explain the paradox. Rather, cancer suppression should be most extreme when the probability of non-cancer death decreases with age (e.g. alligators), maturation is delayed, fecundity rates are low and fecundity increases with age. Thus, the value of cancer suppression is predicted to be lowest in the vole (short lifespan, high fecundity) and highest in the naked mole rat (long lived with late female sexual maturity). The life history of pre-industrial humans likely selected for quite low levels of cancer suppression. In modern humans that live much longer, this level results in unusually high lifetime cancer risks. The model predicts a lifetime risk of 49% compared with the current empirical value of 43%.     

Size,longevity and cancer: age structure

There is significant recent interest in Peto''s paradox and the related problem of the evolution of large, long-lived organisms in terms of cancer robustness. Peto''s paradox refers to the expectation that large, long-lived organisms have a higher lifetime cancer risk, which is not the case: a paradox. This paradox, however, is circular: large, long-lived organisms are large and long-lived because they are cancer robust. Lifetime risk, meanwhile, depends on the age distributions of both cancer and competing risks: if cancer strikes before competing risks, then lifetime risk is high; if not, not. Because no set of competing risks is generally prevalent, it is instructive to temporarily dispose of competing risks and investigate the pure age dynamics of cancer under the multistage model of carcinogenesis. In addition to augmenting earlier results, I show that in terms of cancer-free lifespan large organisms reap greater benefits from an increase in cellular cancer robustness than smaller organisms. Conversely, a higher cellular cancer robustness renders cancer-free lifespan more resilient to an increase in size. This interaction may be an important driver of the evolution of large, cancer-robust organisms.     

Begging and the risk of predation in nestling birds

Theoretical models of the evolution of begging in nestling passerinesassume that begging is costly, either energetically or in termsof predation. However, few empirical measures of these costsexist. We examined whether nestling begging calls could attractpredators to nests by comparing predation rates at artificialnests with and without playbacks of tree swallow begging calls.Nests were baited with quail eggs and placed in pairs on theground or in modified nest-boxes. Nests with playbacks of beggingcalls were depredated before control nests significantly moreoften in both the ground and nest-box trials, suggesting thatpredators may use begging calls to locate nests. These resultssuggest that the risk of nest predation may be increased becauseof calling by nestlings and provide further support for theassumption that conspicuous begging is costly in terms of predation     

Renesting intervals and duration of the incubation and nestling periods of Sprague's Pipits

ABSTRACT.   Sprague's Pipits ( Anthus spragueii ) are one of the least studied birds in North America, with most information about their breeding biology based on anecdotal observations at relatively few nests. I determined the renesting intervals of female Sprague's Pipits in south-central Saskatchewan and also summarized data about incubation and nestling periods of Sprague's Pipits in Saskatchewan and Manitoba to provide more robust estimates of these life-history parameters. Two of eight radio-marked females initiated replacement clutches after failed nesting attempts, with renesting intervals of 11–13 and 15–16 d, respectively. Of 74 banded females, 3 were subsequently recaptured at later nests during the same breeding season. Nests of two of these females failed due to inclement weather and their renesting attempts were initiated 6 and 7–11 d later, respectively. The third female successfully fledged young and then initiated another nest 17 d later. The new nests of females that renested were all located less than 150 m from previous nests. Incubation and nestling periods for known-aged nests ranged from 12 to 15 d (13.4 ± 0.3 [SE], N = 9) and 11 to 14 d (12.1 ± 0.2, N = 39), respectively. Variation in the length of both incubation and nestling periods was likely a function of prolonged periods of inclement weather. More data are required to determine the extent of renesting in different habitats and the extent to which Sprague's Pipits are capable of successfully raising multiple broods in a single breeding season.     

Fair-balance paradox, star-tree paradox, and Bayesian phylogenetics

The star-tree paradox refers to the conjecture that the posterior probabilities for the three unrooted trees for four species (or the three rooted trees for three species if the molecular clock is assumed) do not approach 1/3 when the data are generated using the star tree and when the amount of data approaches infinity. It reflects the more general phenomenon of high and presumably spurious posterior probabilities for trees or clades produced by the Bayesian method of phylogenetic reconstruction, and it is perceived to be a manifestation of the deeper problem of the extreme sensitivity of Bayesian model selection to the prior on parameters. Analysis of the star-tree paradox has been hampered by the intractability of the integrals involved. In this article, I use Laplacian expansion to approximate the posterior probabilities for the three rooted trees for three species using binary characters evolving at a constant rate. The approximation enables calculation of posterior tree probabilities for arbitrarily large data sets. Both theoretical analysis of the analogous fair-coin and fair-balance problems and computer simulation for the tree problem confirmed the existence of the star-tree paradox. When the data size n --> infinity, the posterior tree probabilities do not converge to 1/3 each, but they vary among data sets according to a statistical distribution. This distribution is characterized. Two strategies for resolving the star-tree paradox are explored: (1) a nonzero prior probability for the degenerate star tree and (2) an increasingly informative prior forcing the internal branch length toward zero. Both appear to be effective in resolving the paradox, but the latter is simpler to implement. The posterior tree probabilities are found to be very sensitive to the prior.     

Constitutive immune defences correlate with life-history variables in tropical birds

1. It has been suggested that immune defences are shaped by life history and ecology, but few general patterns have been described across species. We hypothesized that 'fast' life-history traits (e.g. short development times, large clutch sizes) would be associated with developmentally inexpensive immune defences, minimizing the resource demands of young animals' immune systems during periods of rapid growth. Conversely, 'slow' life histories should be associated with well developed antibody-mediated defences, which are developmentally costly. 2. We therefore predicted that 'fast-living' species would exhibit higher levels of complement proteins, a component of non-specific innate defence, but lower levels of constitutive ('natural') antibodies. Additionally, we predicted that constitutive immune defences in general would be higher in species with ecological characteristics that might increase exposure to pathogens, such as open nests, omnivorous diets, gregariousness, and closed forested habitat. 3. Across 70 Neotropical bird species, we found a strongly positive relationship between incubation period and natural antibody levels in adult birds, suggesting that longer developmental times might allow the production of a more diverse and/or more reactive adaptive immune system. Complement activity was positively, although weakly, correlated with clutch size, providing some support for the hypothesis that faster-living species rely more on innate defences, such as complement. Unexpectedly, solitary species had higher natural antibody titres than species that frequently join flocks. 4. Our results suggest that, despite probably widespread differences in the intensity and diversity of pathogen exposure, species-level variation in constitutive immune defences is understandable within the context of life-history theory.     

Breeding biology of Eared Quetzals in the Sierra Madre Occidental, Mexico

ABSTRACT.   Eared Quetzals ( Euptilotis neoxenus ), a threatened species, are one of the least studied trogons in Mexico. We monitored 29 Eared Quetzal nests in the Chihuahuan portion of the Sierra Madre Occidental from 1998 to 2003. All nests were in tree cavities, and the mean tree and nest cavity heights ( N = 14) were 16.9 ± 7.8 m and 11.4 ± 4.1 m, respectively. The mean clutch size was 2.8 ± 0.9 eggs ( N = 28), the incubation period lasted 22 d ( N = 1), and nestling periods ranged from 29 to 31 d ( N = 5). Both adults incubated eggs and fed nestlings. Of 80 eggs, 70 hatched (87.5%) and 67 of 70 young fledged (95.7%). Twenty-five of 29 nests (86.2%) produced at least one fledgling. One nest was predated, and two failed when nest trees fell. Higher rates of nest predation have been reported for other species of trogons. However, fewer potential predators, such as snakes and mammals, are present in the Sierra Madre than in tropical zones where most trogon species occur. In addition, antipredator behaviors, including nestlings with calls resembling a snake and nests with an unpleasant odor, may contribute to the high nesting success. The main limiting factors for Eared Quetzals in the northern Chihuahua may be competition for cavities with other secondary cavity-nesters, and the failure of nests when snags fall.     

Nest attendance by tropical and temperate passerine birds: Same constancy,different strategy

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Life history, ecology and parasite community structure in Soviet birds

The paper describes an investigation of parasite richness in relation to host life history and ecology using data from an extensive survey of helminth parasites (cestodes, trematodes and nematodes) in Soviet birds. Correlates of parasite richness (number of parasite species per host species) were sought among 13 life-history variables, 13 ecological variables and one non-biological variable (number of host individuals examined) across a sample of 158 species of host. A statistical method to control for the effects of phylogenetic association was adopted throughout. Parasite richness correlates positively with the number of hosts examined (sample size) in all three parasite groups. Positive correlations (after controlling for the effects of sample size) were also found between host body weight and parasite richness for trematodes and nematodes, but not for cestodes.
A number of ecological variables were associated with parasite richness. However, when the effects of sample size and body weight were controlled for, only a single significant correlation (an association between trematode richness and aquatic habitat) remained. Similarly, a number of significant correlates of parasite richness were found among the life-history variables examined. Though several of these were robust to the confounding effects of sample size, all could be explained by the co-variation between life-history traits and body weight among the host species under investigation.     

Interspecific variation in the use of carotenoid-based coloration in birds: diet, life history and phylogeny

Birds show striking interspecific variation in their use of carotenoid-based coloration. Theory predicts that the use of carotenoids for coloration is closely associated with the availability of carotenoids in the diet but, although this prediction has been supported in single-species studies and those using small numbers of closely related species, there have been no broad-scale quantitative tests of the link between carotenoid coloration and diet. Here we test for such a link using modern comparative methods, a database on 140 families of birds and two alternative avian phylogenies. We show that carotenoid pigmentation is more common in the bare parts (legs, bill and skin) than in plumage, and that yellow coloration is more common than red. We also show that there is no simple, general association between the availability of carotenoids in the diet and the overall use of carotenoid-based coloration. However, when we look at plumage coloration separately from bare part coloration, we find there is a robust and significant association between diet and plumage coloration, but not between diet and bare part coloration. Similarly, when we look at yellow and red plumage colours separately, we find that the association between diet and coloration is typically stronger for red coloration than it is for yellow coloration. Finally, when we build multivariate models to explain variation in each type of carotenoid-based coloration we find that a variety of life history and ecological factors are associated with different aspects of coloration, with dietary carotenoids only being a consistent significant factor in the case of variation in plumage. All of these results remain qualitatively unchanged irrespective of the phylogeny used in the analyses, although in some cases the precise life history and ecological variables included in the multivariate models do vary. Taken together, these results indicate that the predicted link between carotenoid coloration and diet is idiosyncratic rather than general, being strongest with respect to plumage colours and weakest for bare part coloration. We therefore suggest that, although the carotenoid-based bird plumage may a good model for diet-mediated signalling, the use of carotenoids in bare part pigmentation may have a very different functional basis and may be more strongly influenced by genetic and physiological mechanisms, which currently remain relatively understudied.     

Breeding biology of Orange‐breasted (Harpactes oreskios) and Red‐headed (H. erythrocephalus) trogons in Khao Yai National Park,Thailand

ABSTRACT As tropical habitats continue to be cleared or degraded, obtaining basic information about the ecology of birds in intact habitats is essential for understanding their life histories. We studied the breeding biology of Orange‐breasted Trogons (Harpactes oreskios) and Red‐headed Trogons (H. erythrocephalus) in Khao Yai National Park in Thailand from 2003 to 2009. Nests were in excavated cavities in well‐rotted stumps or other tree parts. Mean cavity heights were 2.1 m (N= 19) for Orange‐breasted Trogons and 2.0 m (N= 49) for Red‐headed Trogons. Eggs were laid every other day. For Orange‐breasted Trogons, the mean clutch size was 2.4 ± 0.1 (SE) eggs (N= 17); incubation periods for two nests were 17 and 18 d, respectively, and the nestling period ranged from 12 to at least 14 d (N= 4). For Red‐headed Trogons, the mean clutch size was 2.6 ± 0.1 eggs (N= 48), the mean incubation period was 18 d (N= 9), and the mean nestling period was 13.4 d (N= 5). In both species, both males and females excavated nest sites, incubated eggs, and brooded and provisioned nestlings. Only females incubated and brooded at night, and males provisioned nestlings more than females. Breeding seasons lasted from January to March for Orange‐breasted Trogons, and from late February to July for Red‐headed Trogons. Mayfield estimates of nest success were 8% and 9% for Orange‐breasted and Red‐headed trogons, respectively. Unusual for cavity nesters, nest failure due to predation was high and nestling periods short. The low nesting success is typical of many other tropical species, but considerably lower than reported for some Neotropical trogons, possibly due to the unenclosed structure of the nests of these Asian trogons.     

Borders,paradox and power

Border studies have grappled with, on the one hand, the need for the use of common themes or concepts while, on the other, the need for contextual specificity. Borders are sites that embody different potentialities: division and contact, conflict and cooperation, security and anxiety, creativity and oppression, among others. In short, they are sites of the paradoxical. Paradox, it is argued, is the common overarching conceptual characteristic of borders but which specific potentialities are embodied in a border and what prevails as a result of the ensuing power struggles requires contextual specificity. Cyprus, a divided island lying on various border lines, partly inside and partly outside the EU, presents a useful socio-political space in order to illustrate this argument by outlining the specific paradoxical aspects of its own border and the results of the ensuing power struggles.     

Temperature,size and developmental plasticity in birds

As temperatures increase, there is growing evidence that species across much of the tree of life are getting smaller. These climate change-driven size reductions are often interpreted as a temporal analogue of the observation that individuals within a species tend to be smaller in the warmer parts of the species'' range. For ectotherms, there has been a broad effort to understand the role of developmental plasticity in temperature–size relationships, but in endotherms, this mechanism has received relatively little attention in favour of selection-based explanations. We review the evidence for a role of developmental plasticity in warming-driven size reductions in birds and highlight insulin-like growth factors as a potential mechanism underlying plastic responses to temperature in endotherms. We find that, as with ectotherms, changes in temperature during development can result in shifts in body size in birds, with size reductions associated with warmer temperatures being the most frequent association. This suggests developmental plasticity may be an important, but largely overlooked, mechanism underlying warming-driven size reductions in endotherms. Plasticity and natural selection have very different constraining forces, thus understanding the mechanism linking temperature and body size in endotherms has broad implications for predicting future impacts of climate change on biodiversity.     

Latitudinal extent and natural history characteristics of birds in Nicaragua

This paper assesses the latitudinal extent of terrestrial breeding birds in Nicaragua. In particular, associations among latitudinal midpoint, body mass, and latitudinal extent are examined; significant differences between natural history characteristics (trophic guild, forest dependence, number of forest types) and latitudinal extent are identified; and a test is undertaken of Rapoport's rule for birds at the edge of their northern or southern range in Nicaragua. Birds in Nicaragua were classified into four categories based on latitudinal extent: birds generally restricted to Central America (20%); birds near the edge of their range within Nicaragua (34%); Neotropical birds (42%); and Pan‐American birds (4%). Latitudinal midpoint had a significant negative correlation with latitudinal extent over different taxonomic scales. Natural history characteristics of trophic guild, forest dependence and number of forest types can account for significant differences in species latitudinal extent. Carnivores had greater latitudinal extents than most other trophic guilds, non‐forest birds had greater latitudinal extents than birds that require patchy forest, and birds that occur in four or more forest types had greater latitudinal extents than birds restricted to one or two forest types. Contrary to Rapoport's rule, birds with a northern affinity or latitudinal midpoint north of Nicaragua had significantly smaller latitudinal extents than birds with a southern affinity or latitudinal midpoint south of Nicaragua. A comparison of natural history characteristics of birds with northern and southern affinities found no difference between trophic guild or forest dependence but a significant difference among the number of forest types used by forest birds. Birds with a southern affinity were restricted to fewer forest types than birds with a northern affinity.     

Life history and the evolution of family living in birds

The reason why some bird species live in family groups is an important question of evolutionary biology that remains unanswered. Families arise when young delay the onset of independent reproduction and remain with their parents beyond independence. Explanations for why individuals forgo independent reproduction have hitherto focused on dispersal constraints, such as the absence of high-quality breeding openings. However, while constraints successfully explain within-population dispersal decisions, they fail as an ultimate explanation for variation in family formation across species. Most family-living species are long-lived and recent life-history studies demonstrated that a delayed onset of reproduction can be adaptive in long-lived species. Hence, delayed dispersal and reproduction might be an adaptive life-history decision rather than 'the best of a bad job'. Here, we attempt to provide a predictive framework for the evolution of families by integrating life-history theory into family formation theory. We suggest that longevity favours a delayed onset of reproduction and gives parents the opportunity of a prolonged investment in offspring, an option which is not available for short-lived species. Yet, parents should only prolong their investment in offspring if this increases offspring survival and outweighs the fitness cost that parents incur, which is only possible under ecological conditions, such as a predictable access to resources. We therefore propose that both life-history and ecological factors play a role in determining the evolution of family living across species, yet we suggest different mechanisms than those proposed by previous models.     

Multiple responses to increasing spring temperatures in the breeding cycle of blue and great tits (Cyanistes caeruleus,Parus major)

Many bird species start laying their eggs earlier in response to increasing spring temperatures, but the causes of variation between and within species have not been fully explained. Moreover, synchronization of the nestling period with the food supply not only depends on first‐egg dates but also on additional reproductive parameters including laying interruptions, incubation time and nestling growth rate. We studied the breeding cycle of two sympatric and closely related species, the blue tit Cyanistes caeruleus and the great tit Parus major in a rich oak‐beech forest, and found that both advanced their mean first‐egg dates by 11–12 days over the last three decades. In addition, the time from first egg to fledging has shortened by 2–3 days, through a decrease in laying interruptions, incubation time (not statistically significant) and nestling development time. This decrease is correlated with a gradual increase of temperatures during laying, suggesting a major effect of the reduction in laying interruptions. In both species, the occurrence of second clutches has strongly decreased over time. As a consequence, the average time of fledging (all broods combined) has advanced by 15.4 and 18.6 days for blue and great tits, respectively, and variance in fledging dates has decreased by 70–75%. Indirect estimates of the food peak suggest that both species have maintained synchronization with the food supply. We found consistent selection for large clutch size, early laying and short nest time (laying to fledging), but no consistent changes in selection over time. Analyses of within‐individual variation show that most of the change can be explained by individual plasticity in laying date, fledging date and nest time. This study highlights the importance of studying all components of the reproductive cycle, including second clutches, in order to assess how natural populations respond to climate change.     

Egg mass and incubation period allometry in birds and reptiles: effects of phylogeny

Here we test the hypothesis that the relationship between egg mass at oviposition (IEM) and incubation period ( I _p) is a function of the taxonomic relatedness of bird and reptile species. Allometric relationships between IEM and I _p were examined for 1525 bird species and 201 reptilian species. Treating species as independent data revealed the allometric exponent linking I _p to IEM to be 0.234 for birds and 0.138 for reptiles. However, ANCOVA revealed that within both birds and reptiles the elevation and slope of the regression lines were dependent on the taxonomic order studied, indicating that the exponents were confounded by the phylogenetic relatedness of species. Thus, allometric exponents were recalculated based on the method of comparative analysis using independent contrasts. This technique revealed that the allometric exponent in both birds and reptiles was confounded by phylogeny. In birds the allometric relationship between I _p and IEM was almost halved to 0.122, whereas in reptiles the exponent increased to 0.185. Importantly, the results demonstrate that some results of allometric analyses can be artefacts of the method of analysis of the dataset. That for bird eggs I _p is not determined in large part by egg mass allows new questions to be posed regarding the ecological and physiological factors affecting the length of incubation, and hence rates of embryonic growth, for different taxa and habitats.     

Life history patterns in birds and mammals and their evolutionary interpretation

Summary It is argued that allometric principles account for most of the observed variation in the life history patterns amongst birds. To test this contention it is shown that traits such as incubation time, growth rates, age at first reproduction, lifespan, clutch weight and egg weight all scale to body weight with exponents similar to those found for analogous traits in mammals. It is then shown that most of the variation amongst bird taxa and between birds and mammals based on body weight allometry can be explained by variations in brain size, body temperature and metabolic rate, consistent with theories of growth and ageing derived from mammalian studies. Finally, it is suggested that the evidence for life histostory allometry is sufficiently strong that it argues for a more epigenetic view of life history patterns and their evolution than is generally conceded in most adaptation theories.