Phyletic size change and brain/body allometry: A consideration based on the African pongids and other primates

A problematic aspect of brain/body allometry is the frequency of interspecific series which exhibit allometry coefficients of approximately 0.33. This coefficient is significantly lower than the 0.66 value which is usually taken to be the interspecific norm. A number of explanations have been forwarded to account for this finding. These include (1) intraspecificallometry explanations, (2) nonallometric explanations, and (3) Jerison’s “extraneurons” hypothesis, among others. The African apes, which exhibit a lowered interspecific allometry coefficient, are used here to consider previous explanations. These are found to be inadequate in a number of ways, and an alternative explanation is proposed. This explanation is based on patterns of brain and body size change during ontogeny and phytogeny. It is argued that the interspecific allometry coefficient in African apes parallels the intraspecific one because similar ontogenetic modifications of body growth separate large and small forms along each curve. In both cases, body size differences are produced primarily by growth in later postnatal periods, during which little brain growth occurs. Data on body growth, neonatal scaling, and various lifehistory traits support this explanation. This work extends previous warnings that sizecorrected estimates of relative brain size may not correspond very closely to our understanding of the behavioral capacities of certain species in lineages characterized by rapid change in body size.     

Testing the cranial evolutionary allometric ‘rule’ in Galliformes

Recent comparative studies have indicated the existence of a common cranial evolutionary allometric (CREA) pattern in mammals and birds, in which smaller species have relatively smaller faces and bigger braincases than larger species. In these studies, cranial allometry was tested using a multivariate regression between shape (described using landmarks coordinates) and size (i.e. centroid size), after accounting for phylogenetic relatedness. Alternatively, cranial allometry can be determined by comparing the sizes of two anatomical parts using a bivariate regression analysis. In this analysis, a slope higher or lower than one indicates the existence of positive or negative allometry, respectively. Thus, in those species that support the CREA ‘rule’, positive allometry is expected for the association between face size and braincase size, which would indicate that larger species have disproportionally larger faces. In this study, I applied these two approaches to explore cranial allometry in 83 Galliformes (Aves, Galloanserae), ranging in mean body weight from 30 g to 2.5 kg. The multivariate regression between shape and centroid size revealed the existence of a significant allometric pattern resembling CREA, whereas the second analysis revealed a negative allometry for beak size and braincase size (i.e. contrary to the CREA ‘rule’, larger galliform species have disproportionally shorter beaks than smaller galliform species). This study suggests that the presence of CREA may be overestimated when using cranium size as the standard measurement.     

Variation across amphibian species in the size of the nuclear genome supports a pluralistic, hierarchical approach to the C-value enigma

Amphibians have featured prominently in discussions of the C-value enigma, the still-unresolved puzzle regarding the evolution of genome size. Their wide range in nuclear DNA contents and diverse ecological and developmental lifestyles make them excellent subjects for addressing the key elements of the C-value enigma. However, in some cases the importance of work on amphibians appears to be overstated. This is especially true of claims that patterns of variation in salamanders support a particular theory of genome size evolution to the exclusion of others. This study provides a critical re-examination of some of these claims, as well as an investigation of the relationships between genome size, cell and nuclear size, and metabolism in amphibians. The results of these analyses, combined with an overview of previous amphibian genome size literature, strongly indicate the need for a pluralistic approach to the C-value enigma. In particular, it must be recognized that evolutionary forces operating and interacting at several levels of biological organization (of which the genome itself is one) are responsible for the observed patterns in amphibian genome size distributions.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 329–339.     

Patterns of sexual size dimorphism in Chelonia

Macroevolutionary patterns of sexual size dimorphism (SSD) indicate how sexual selection, natural selection, and genetic and developmental constraints mold sex differences in body size. One putative pattern, known as Rensch's rule, posits that, among species with female‐larger SSD, the relative degree of SSD declines with species' body size, whereas, among male‐larger SSD species, relative SSD increases with size. Using a dataset of 196 chelonian species from all fourteen families, we investigated the correlation in body size evolution between male and female Chelonia and the validity of Rensch's rule for the taxon and within its major clades. We conclude that male–female correlations in body size evolution are high, although these correlations differ among chelonian families. Overall, SSD scales isometrically with body size; Rensch's rule is valid for only one family, Testudinidae (tortoises). Because macroevolutionary patterns of SSD can vary markedly among clades, even in a taxon as morphologically conservative as Testudines, one must guard against inappropriately pooling clades in comparative studies of SSD. The results of the present study also indicate that regression models that assume the x‐variable (e.g. male body size) is measured without statistical error, although frequently reported, will result in erroneous conclusions about phylogenetic trends in sexual size dimorphism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 396–413.     

Large brain size is associated with low extra‐pair paternity across bird species

BackgroundGaining extrapair copulations (EPCs) is a complicated behavior process. The interaction between males and females to procure EPCs may be involved in brain function evolution and lead to a larger brain. Thus, we hypothesized that extrapair paternity (EPP) rate can be predicted by relative brain size in birds. Past work has implied that the EPP rate is associated with brain size, but empirical evidence is rare.MethodsWe collated data from published references on EPP levels and brain size of 215 bird species to examine whether the evolution of EPP rate can be predicted by brain size using phylogenetically generalized least square (PGLS) models and phylogenetic path analyses.ResultsWe found that EPP rates (both the percentage EP offspring and percentage of broods with EP offspring) are negatively associated with relative brain size. We applied phylogenetic path analysis to test the causal relationship between relative brain size and EPP rate. Best‐supported models (ΔCICc < 2) suggested that large brain lead to reduced EPP rate, which failed to support the hypothesis that high rates of EPP cause the evolution of larger brains.ConclusionThis study indicates that pursuing EPCs may be a natural instinct in birds and the interaction between males and females for EPCs may lead to large brains, which in turn may restrict their EPC level for both sexes across bird species.     

Acclimation of respiration to temperature and CO2 in seedlings of boreal tree species in relation to plant size and relative growth rate

The role of acclimation of dark respiration to temperature and CO₂ concentration and its relationship to growth are critical in determining plant response to predicted global change. We explored temperature acclimation of respiration in seedlings of tree species of the North American boreal forest. Populus tremuloides, Betula papyrifera, Larix laricina, Pinus banksiana, and Picea mariana plants were grown from seed in controlled-environments at current and elevated concentrations of CO₂ (370 and 580 μmol mol^–1) in combination with three temperature treatments of 18/12, 24/18, and 30/24 °C (light/dark period). Specific respiration rates of roots and shoots acclimated to temperature, damping increases in rates across growth-temperature environments compared to short-term temperature responses. Compared at a standard temperature, root and shoot respiration rates were, on average, 40% lower in plants grown at the highest compared to lowest growth temperature. Broad-leaved species had a lower degree of temperature acclimation of respiration than did the conifers. Among species and treatment combinations, rates of respiration were linearly related to size and relative growth rate, and relationships were comparable among growth environments. Specific respiration rates and whole-plant respiratory CO₂ efflux as a proportion of daily net CO₂ uptake increased at higher growth temperatures, but were minimally affected by CO₂ concentration. Whole-plant specific respiration rates were two to three times higher in broad-leaved than coniferous species. However, compared to faster-growing broad-leaved species, slower-growing conifers lost a larger proportion of net daily CO₂ uptake as respiratory CO₂ efflux, especially in roots. Interspecific variation in acclimation responses of dark respiration to temperature is more important than acclimation of respiration to CO₂ enrichment in modifying tree seedling growth responses to projected increases in CO₂ concentration and temperature.     

异速生长模型研究概述

最近,关于异速生长模型的讨论再次成为焦点,讨论热点为异速生长指数的取值及其理论解释.本文综述了WBE 97、BMR(99)模型的相关研究,重点介绍了MGL模型及由此模型得到的结果:个体整体的新陈代谢率与个体的质量没有明显依赖关系,其标度指数不是一个固定的值,而是一个区间[2/3,1].考虑的视角从个体整体的新陈代谢率转到单位质量的新陈代谢率,通过对不同物种、不同环境的单位质量新陈代谢率的研究,发现对大多数物种,其值落在一个具有普适性的上、下界的区间内;认为存在单位质量的新陈代谢率最小值确定了个体的大小,并建立基于该最小值的描述个体大小与温度关系的数学模型,该模型得到实验数据验证.     

Ontogenic changes in the allometric scaling of the mass and length relationship in Sprattus sprattus

An analysis of mass ( M ) and standard length ( L _S) data for larval, juvenile and adult sprat ( Sprattus sprattus ; Clupeidae) revealed marked changes in the allometric scaling factor ( b in     ). For sprat <44 mm L _S, b was 5·0, whereas in larger juveniles and adults, b was c. 3·4 indicating a relatively protracted metamorphic period for this species.     

Early development and allometric growth patterns in Siberian sturgeon and their ecological significance

Early development of Siberian sturgeon Acipenser baeri was divided into two different phases, the prelarval stage between hatching (10·4–11·1 mm L_T) and first feeding (19·6–21·0 mm L_T), and the larval stage between the initiation of external feeding and metamorphosis (28·6–32·4 mm L_T). Morphogenesis and differentiation were more intense during the prelarval than larval and early juvenile stages; the prelarval period was characterized by the replacement of embryonic adaptations and functions by definitive ones, such as branchial respiration, exogenous feeding, and active swimming. The positive allometry of the head for feeding, sensorial and respiratory functions (inflexion point at 20·0 mm L_T), and the tail for reducing costs of transport, routine swimming and escape reactions from predators (inflexion point at 20·2 mm L_T) confirmed the hypothesis that growth patterns of early life stages closely match specific needs.     

The evolution of hippocampus volume and brain size in relation to food hoarding in birds

Food‐hoarding birds frequently use spatial memory to relocate their caches, thus they may evolve a larger hippocampus in their brain than non‐hoarder species. However, previous studies testing for such interspecific relationships provided conflicting results. In addition, food hoarding may be a cognitively complex task involving elaboration of a variety of brain regions, even outside of the hippocampus. Hence, specialization to food hoarding may also result in the enlargement of the overall brain. In a phylogenetic analysis of distantly related birds, we studied the interspecific association between food hoarding and the size of different brain regions, each reflecting different resolutions. After adjusting for allometric effects, the relative volume of the hippocampus and the relative size of the entire brain were each positively related to the degree of food‐hoarding specialization, even after controlling for migration and brood parasitism. We also found some significant evidence for the relative volume of the telencephalon being associated with food hoarding, but this relationship was dependent on the approach we used. Hence, neural adaptation to food hoarding may favour the evolution of different brain structures.     

The allometric pattern of sexually size dimorphic feather ornaments and factors affecting allometry

The static allometry of secondary sexual characters is currently subject to debate. While some studies suggest an almost universal positive allometry for such traits, but isometry or negative allometry for nonornamental traits, other studies maintain that any kind of allometric pattern is possible. Therefore, we investigated the allometry of sexually size dimorphic feather ornaments in 67 species of birds. We also studied the allometry of female feathers homologous to male ornaments (female ornaments in the following) and ordinary nonsexual traits. Allometries were estimated as reduced major axis slopes of trait length on tarsus length. Ornamental feathers showed positive allometric slopes in both sexes, although that was not a peculiarity for ornamental feathers, because nonsexual tail feathers also showed positive allometry. Migration distance (in males) and relative size of the tail ornament (in females) tended to be negatively related to the allometric slope of tail feather ornaments, although these results were not conclusive. Finally, we found an association between mating system and allometry of tail feather ornaments, with species with more intense sexual selection showing a smaller degree of allometry of tail ornaments. This study is consistent with theoretical models that predict no specific kind of allometric pattern for sexual and nonsexual characters.     

A comparative perspective on longevity: the effect of body size dominates over ecology in moths

Both physiologically and ecologically based explanations have been proposed to account for among‐species differences in lifespan, but they remain poorly tested. Phylogenetically explicit comparative analyses are still scarce and those that exist are biased towards homoeothermic vertebrates. Insect studies can significantly contribute as lifespan can feasibly be measured in a high number of species, and the selective forces that have shaped it may differ largely between species and from those acting on larger animals. We recorded adult lifespan in 98 species of geometrid moths. Phylogenetic comparative analyses were applied to study variation in species‐specific values of lifespan and to reveal its ecological and life‐history correlates. Among‐species and between‐gender differences in lifespan were found to be notably limited; there was also no evidence of phylogenetic signal in this trait. Larger moth species were found to live longer, with this result supporting a physiological rather than ecological explanation of this relationship. Species‐specific lifespan values could not be explained by traits such as reproductive season and larval diet breadth, strengthening the evidence for the dominance of physiological determinants of longevity over ecological ones.     

Body size, egg size, and their interspecific relationships with ecological and life history traits in butterflies (Lepidoptera: Papilionoidea, Hesperioidea)

The interspecific relationships between egg size and body size in butterflies (Papilionoidea and Hesperiidae), and between size and egg and larval development time, larval trophic specificity, foodplant structure, climate, and phenology were investigated based on a sample of more than 1180 species. The independent contrasts mediod was used to avoid taxonomy-dependent results. Egg size is allometrically related to adult wing length by a slope of 0.43. Based on a subset of species, fecundity is correlated to adult body size, and there is evidence for a compromise between egg number and egg size (relative to adult size) across species. Butterfly size increases in correlation to the mean annual temperature of me species geographic range, but decreases in relation to increased aridity (or die length of the dry season). Larger butterflies tend to have longer larval development times, use large or structurally complex host plants, and are more likely to lay their eggs in batches, irrespective of climate. Larger eggs tend to develop more slowly, and give rise to larvae with longer developmental periods that will result in larger adults. No evidence was found to support a relationship between butterfly body size and polyphagy. A complex pattern of interrelationships links body size (and egg size) to other traits, although correlations other than mat between egg size and body size are generally low. The results suggest the necessity of separating climate and seasonality into components that are relevant to insect life histories in comparative studies.     

The evolution of brain size and intelligence in man

The brain size of hominids has increased approximately threefold during the evolution of the hominids fromAustralopithecus toHomo sapiens. It is proposed that the principal reason for this increase is that larger brains conferred greater intelligence, and greater intelligence conferred a selection advantage. A number of anthropologists have difficulty accepting this thesis because they believe that brain size is not associated with intelligence in man. Evidence is reviewed, and new evidence from two studies is presented, to show that brain size as measured by head size is positively correlated with intelligence as measured by intelligence tests. On two recent samples statistically significant correlations of .21 and .30 were obtained between estimates of brain size and IQ. It is considered that brain size is positively associated with intelligence in man and that this is the major reason for the increase in brain size of the hominids during the last 3.2 million years.     

Gill surface area provides a clue for the respiratory basis of brain size in the blacktip shark (Carcharhinus limbatus)

Brain size varies dramatically, both within and across species, and this variation is often believed to be the result of trade-offs between the cognitive benefits of having a large brain for a given body size and the energetic cost of sustaining neural tissue. One potential consequence of having a large brain is that organisms must also meet the associated high energetic demands. Thus, a key question is whether metabolic rate correlates with brain size. However, using metabolic rate to measure energetic demand yields a relatively instantaneous and dynamic measure of energy turnover, which is incompatible with the longer evolutionary timescale of changes in brain size within and across species. Morphological traits associated with oxygen consumption, specifically gill surface area, have been shown to be correlates of oxygen demand and energy use, and thus may serve as integrated correlates of these processes, allowing us to assess whether evolutionary changes in brain size correlate with changes in longer-term oxygen demand and energy use. We tested how brain size relates to gill surface area in the blacktip shark Carcharhinus limbatus. First, we examined whether the allometric slope of brain mass (i.e., the rate that brain mass changes with body mass) is lower than the allometric slope of gill surface area across ontogeny. Second, we tested whether gill surface area explains variation in brain mass, after accounting for the effects of body mass on brain mass. We found that brain mass and gill surface area both had positive allometric slopes, with larger individuals having both larger brains and larger gill surface areas compared to smaller individuals. However, the allometric slope of brain mass was lower than the allometric slope of gill surface area, consistent with our prediction that the allometric slope of gill surface area could pose an upper limit to the allometric slope of brain mass. Finally, after accounting for body mass, individuals with larger brains tended to have larger gill surface areas. Together, our results provide clues as to how fishes may evolve and maintain large brains despite their high energetic cost, suggesting that C. limbatus individuals with a large gill surface area for their body mass may be able to support a higher energetic turnover, and, in turn, a larger brain for their body mass.     

Lowering sample size in comparative analyses can indicate a correlation where there is none: example from Rensch's rule in primates

The fact that characters may co-vary in organism groups because of shared ancestry and not always because of functional correlations was the initial rationale for developing phylogenetic comparative methods. Here we point out a case where similarity due to shared ancestry can produce an undesired effect when conducting an independent contrasts analysis. Under special circumstances, using a low sample size will produce results indicating an evolutionary correlation between characters where an analysis of the same pattern utilizing a larger sample size will show that this correlation does not exist. This is the opposite effect of increased sample size to that expected; normally an increased sample size increases the chance of finding a correlation. The situation where the problem occurs is when co-variation between the two continuous characters analysed is clumped in clades; e.g. when some phylogenetically conservative factors affect both characters simultaneously. In such a case, the correlation between the two characters becomes contingent on the number of clades sharing this conservative factor that are included in the analysis, in relation to the number of species contained within these clades. Removing species scattered evenly over the phylogeny will in this case remove the exact variation that diffuses the evolutionary correlation between the two characters - the variation contained within the clades sharing the conservative factor. We exemplify this problem by discussing a parallel in nature where the described problem may be of importance. This concerns the question of the presence or absence of Rensch's rule in primates.     

Reproductive strategies,body size,and encephalization in primate evolution

In order to understand fully the generally high level of encephalization observed in living primates, we must determine why early primates exhibited moderately large relative brain sizes compared to their early Tertiary contemporaries. The relatively high degree of encephalization in early primates may be related at least in part to the fact that they were highly unusualamong mammals in combining a small body size with a strongly precocial reporductive strategy. Other small, precocial mammals also exhibit moderately high levels of encephalization; but primates appear to have been truly uniquein being the only such small-sized and highly precocial group to give rise to extensive radiations of larger descendants. This is a key element in understanding primate brain evolution, because the initial “head start” of the early primates was translated up to larger sizes in descendants. The possible relationships among encephalization, precociality, small size, and arboreality are discussed, particularly in light of recent debates concerning the validity of the superorder Archonta. This work emphasizes that we need to consider relative brain size as but one element in a complex synergistic network of morphological and life-history features.     

The influence of size on body shape diversification across Indo‐Pacific shore fishes*

Understanding the causes of body shape variability across the tree of life is one of the central issues surrounding the origins of biodiversity. One potential mechanism driving observed patterns of shape disparity is a strongly conserved relationship between size and shape. Conserved allometry has been shown to account for as much as 80% of shape variation in some vertebrate groups. Here, we quantify the amount of body shape disparity attributable to changes in body size across nearly 800 species of Indo‐Pacific shore fishes using a phylogenetic framework to analyze 17 geometric landmarks positioned to capture general body shape and functionally significant features. In marked contrast to other vertebrate lineages, we find that changes in body size only explain 2.9% of the body shape variation across fishes, ranging from 3% to 50% within our 11 sampled families. We also find a slight but significant trend of decreasing rates of shape evolution with increasing size. Our results suggest that the influence of size on fish shape has largely been overwhelmed by lineage‐specific patterns of diversification that have produced the modern landscape of highly diverse forms that we currently observe in nature.