Models projecting the fate of fish populations under climate change need to be based on valid physiological mechanisms

Some recent modelling papers projecting smaller fish sizes and catches in a warmer future are based on erroneous assumptions regarding (i) the scaling of gills with body mass and (ii) the energetic cost of ‘maintenance’. Assumption (i) posits that insurmountable geometric constraints prevent respiratory surface areas from growing as fast as body volume. It is argued that these constraints explain allometric scaling of energy metabolism, whereby larger fishes have relatively lower mass‐specific metabolic rates. Assumption (ii) concludes that when fishes reach a certain size, basal oxygen demands will not be met, because of assumption (i). We here demonstrate unequivocally, by applying accepted physiological principles with reference to the existing literature, that these assumptions are not valid. Gills are folded surfaces, where the scaling of surface area to volume is not constrained by spherical geometry. The gill surface area can, in fact, increase linearly in proportion to gill volume and body mass. We cite the large body of evidence demonstrating that respiratory surface areas in fishes reflect metabolic needs, not vice versa, which explains the large interspecific variation in scaling of gill surface areas. Finally, we point out that future studies basing their predictions on models should incorporate factors for scaling of metabolic rate and for temperature effects on metabolism, which agree with measured values, and should account for interspecific variation in scaling and temperature effects. It is possible that some fishes will become smaller in the future, but to make reliable predictions the underlying mechanisms need to be identified and sought elsewhere than in geometric constraints on gill surface area. Furthermore, to ensure that useful information is conveyed to the public and policymakers about the possible effects of climate change, it is necessary to improve communication and congruity between fish physiologists and fisheries scientists.     

The allometric approach to species differences in brain size

Allometric methods can be used to test quantitative theories of the relationship between brain size and body size across species, and to search for ecological, behavioural, life history, and ontogenetic correlates of brain size. Brain size scales with an allometric exponent of around 0.75 against body size across mammals, but is closer to 0.56 for birds and for reptiles. The slope of the allometric line often varies depending upon the taxonomic level of analysis. However, this phenomenon, at least in mammals, may be a statistical artifact. Brain size for a given body size (relative brain size) varies among orders in birds and mammals, and some dietary associations with relative brain size have been found in particular taxa. Developmental status at birth is the most consistent correlate of relative brain size: precocial neonates have larger brains for a given maternal size than altricial neonates in both birds and mammals. Altricial neonates, however, have more brain growth following birth, and in birds also have larger relative adult brain sizes. Energetic explanations for differences in neonatal brain growth, although attractive on theoretical grounds, have largely failed to stand up to empirical tests.     

Sexual selection uncouples the evolution of brain and body size in pinnipeds

The size of the vertebrate brain is shaped by a variety of selective forces. Although larger brains (correcting for body size) are thought to confer fitness advantages, energetic limitations of this costly organ may lead to trade-offs, for example as recently suggested between sexual traits and neural tissue. Here, we examine the patterns of selection on male and female brain size in pinnipeds, a group where the strength of sexual selection differs markedly among species and between the sexes. Relative brain size was negatively associated with the intensity of sexual selection in males but not females. However, analyses of the rates of body and brain size evolution showed that this apparent trade-off between sexual selection and brain mass is driven by selection for increasing body mass rather than by an actual reduction in male brain size. Our results suggest that sexual selection has important effects on the allometric relationships of neural development.     

The ecology and macroecology of mammalian home range area

Although many studies employ allometric relationships to demonstrate possible dependence of various traits on body mass, the relationship between home range size and body mass has been perhaps the most difficult to understand. Early studies demonstrated that carnivorous species had larger home ranges than herbivorous species of similar mass. These studies also argued that scaling relations (e.g., slopes) of the former were steeper than those of the latter and explained this in terms of the distribution of food resources, which are more uniformly distributed for most herbivores than for carnivores. In contrast to these studies, we show that scaling relations of home ranges for carnivorous mammals do not differ significantly from those of herbivorous and omnivorous species and that all three exhibit slopes that are significantly steeper than predicted on the basis of energetic requirements. We also demonstrate that home range size is constrained to fit within a polygonal constraint space bounded by lines representing energetic and/or biophysical limitations, which suggests that the log-linear relationship between home range area and mass may not be the appropriate function to compare against the energetically predicted slopes of 0.75 or 1.0. It remains unclear, however, why the slope of the relationship between home range area and body mass, whether based on raw data or on constraint lines, always exceeds that predicted by the energetic needs hypothesis.     

Allometric scaling in small colonies of the scleractinian coral Siderastrea siderea (Ellis and Solander)

Although most physiological traits scale allometrically in unitary organisms, it has been hypothesized that modularity allows for isometric scaling in colonial modular taxa. Isometry would allow increases in size without functional constraints, and is thought to be of central importance to the success of a modular design. Yet, despite its potential importance, scaling in these organisms has received little attention. To determine whether scleractinian corals are free of allometric constraints, we quantified metabolic scaling, measured as aerobic respiration, in small colonies (< or =40 mm in diam.) of the scleractinian Siderastrea siderea. We also quantified the scaling of colony surface area with biomass, since the proposed isometry is contingent upon maintaining a constant ratio of surface area to biomass (or volume) with size. Contrary to the predicted isometry, aerobic respiration scaled allometrically on biomass with a slope (b) of 0.176, and colony surface area scaled allometrically on biomass with a slope of 0.730. These findings indicate that small colonies of S. siderea have disproportionately high metabolic rates and SA:B ratios compared to their larger counterparts. The most probable explanations for the allometric scaling of aerobic respiration are (1) a decline in the SA:B ratio with size such that more surface area is available per unit of biomass for mass transfer in the smallest colonies, and (2) the small size, young age, and disproportionately high growth rates of the corals examined. This allometric scaling also demonstrates that modularity, alone, does not allow small colonies of S. siderea to overcome allometric constraints. Further studies are required to determine whether allometric scaling is characteristic of the full size range of colonies of S. siderea.     

Characterizing morphological (co)variation using structural equation models: Body size,allometric relationships and evolvability in a house sparrow metapopulation

Body size plays a key role in the ecology and evolution of all organisms. Therefore, quantifying the sources of morphological (co)variation, dependent and independent of body size, is of key importance when trying to understand and predict responses to selection. We combine structural equation modeling with quantitative genetics analyses to study morphological (co)variation in a meta‐population of house sparrows (Passer domesticus). As expected, we found evidence of a latent variable “body size,” causing genetic and environmental covariation between morphological traits. Estimates of conditional evolvability show that allometric relationships constrain the independent evolution of house sparrow morphology. We also found spatial differences in general body size and its allometric relationships. On islands where birds are more dispersive and mobile, individuals were smaller and had proportionally longer wings for their body size. Although on islands where sparrows are more sedentary and nest in dense colonies, individuals were larger and had proportionally longer tarsi for their body size. We corroborated these results using simulations and show that our analyses produce unbiased allometric slope estimates. This study highlights that in the short term allometric relationships may constrain phenotypic evolution, but that in the long term selection pressures can also shape allometric relationships.     

Primate brains,the ‘island rule’ and the evolution of Homo floresiensis

The taxonomic status of the small bodied hominin, Homo floresiensis, remains controversial. One contentious aspect of the debate concerns the small brain size estimated for specimen LB1 (Liang Bua 1). Based on intraspecific mammalian allometric relationships between brain and body size, it has been argued that the brain of LB1 is too small for its body mass and is therefore likely to be pathological. The relevance and general applicability of these scaling rules has, however, been challenged, and it is not known whether highly encephalized primates adapt to insular habitats in a consistent manner. Here, an analysis of brain and body size evolution in seven extant insular primates reveals that although insular primates follow the ‘island rule’, having consistently reduced body masses compared with their mainland relatives, neither brain mass nor relative brain size follow similar patterns, contrary to expectations that energetic constraints will favour decreased relative brain size. Brain:body scaling relationships previously used to assess the plausibility of dwarfism in H. floresiensis tend to underestimate body masses of insular primates. In contrast, under a number of phylogenetic scenarios, the evolution of brain and body mass in H. floresiensis is consistent with patterns observed in other insular primates.     

What Makes Vocalisation Frequency an Unreliable Signal of Body Size in Birds? A Study on Black Swans

In many animal species, the frequency (pitch) of vocalisations correlates negatively with body size and may thus signal competitive ability. However, this relationship is absent in other species. Understanding why this difference exists across species may help to explain some of the diversity of vocal communication systems. We assessed whether vocalisation frequency signals body size in black swans (Cygnus atratus), and how this is affected by (i) variation in frequency within individuals and (ii) size variation across individuals. Frequency was correlated with body size and mass, with slopes close to the allometry expected if the birds were maximising sound radiation, but the explained variation in frequency was low. Within‐individual variation in vocalisation frequency was greater in male than female swans, and the reliability of frequency as a signal of size in males was correspondingly lower. A review of the literature on the relationship between the frequency of avian vocalisations and body size also showed smaller effect sizes for more variable vocalisations (birdsongs), than for simpler vocalisations. Vocalisation frequency was more reliably correlated with body size when the sexes were pooled (creating a larger range of variation in size) than when the relationship was examined for either sex separately, although male and female data followed the same allometric line. These results show that variation in frequency within individuals and low variation in size across individuals reduce the reliability of vocalisation frequency as a signal of body size, which helps to understand differences among species in the signal value of vocalisation frequency.     

Gregariousness increases brain size in ungulates

The brain’s main function is to organise the physiological and behavioural responses to environmental and social challenges in order to keep the organism alive. Here, we studied the effects that gregariousness (as a measurement of sociality), dietary habits, gestation length and sex have on brain size of extant ungulates. The analysis controlled for the effects of phylogeny and for random variability implicit in the data set. We tested the following groups of hypotheses: (1) Social brain hypothesis—gregarious species are more likely to have larger brains than non-gregarious species because the former are subjected to demanding and complex social interactions; (2) Ecological hypothesis—dietary habits impose challenging cognitive tasks associated with finding and manipulating food (foraging strategy); (3) Developmental hypotheses (a) energy strategy: selection for larger brains operates, primarily, on maternal metabolic turnover (i.e. gestation length) in relation to food quality because the majority of the brain’s growth takes place in utero, and finally (b) sex hypothesis: females are expected to have larger brains than males, relative to body size, because of the differential growth rates of the soma and brain between the sexes. We found that, after adjusting for body mass, gregariousness and gestation length explained most of the variation in brain mass across the ungulate species studied. Larger species had larger brains; gregarious species and those with longer gestation lengths, relative to body mass, had larger brains than non-gregarious species and those with shorter gestation lengths. The effect of diet was negligible and subrogated by gestation length, and sex had no significant effect on brain size. The ultimate cause that could have triggered the co-evolution between gestation length and brain size remains unclear.     

Evolution of brain region volumes during artificial selection for relative brain size

The vertebrate brain shows an extremely conserved layout across taxa. Still, the relative sizes of separate brain regions vary markedly between species. One interesting pattern is that larger brains seem associated with increased relative sizes only of certain brain regions, for instance telencephalon and cerebellum. Till now, the evolutionary association between separate brain regions and overall brain size is based on comparative evidence and remains experimentally untested. Here, we test the evolutionary response of brain regions to directional selection on brain size in guppies (Poecilia reticulata) selected for large and small relative brain size. In these animals, artificial selection led to a fast response in relative brain size, while body size remained unchanged. We use microcomputer tomography to investigate how the volumes of 11 main brain regions respond to selection for larger versus smaller brains. We found no differences in relative brain region volumes between large‐ and small‐brained animals and only minor sex‐specific variation. Also, selection did not change allometric scaling between brain and brain region sizes. Our results suggest that brain regions respond similarly to strong directional selection on relative brain size, which indicates that brain anatomy variation in contemporary species most likely stem from direct selection on key regions.     

Allometry of gill dimensions in some British and American decapod crustacea

The gill areas and their component measurements of 16 species of North American crab and seven species of British decapod crustaceans have been analysed in relation to body mass using the method of logarithmic transformation.
A wide range of relationships was found, each of which is typical for a given species. The slope, b , of the log/log regression lines varied from 0·5 to 1·0, the lower values being most commonly found in the Macrura. For the North American species the average slope is about 0·8 whereas for British brachyurans the relationship was close to linear (b=0·97).
The analysis shows that the increase in gill surface body size is mainly due to the increasing area for individual platelets or gill lamellae.
Comparison of weight-specific gill areas for animals of the same body weight suggests that the most active species have larger gill areas. For some of these species the values (900mm²/g) approximate to those of active fish.
As plots for interspecific relationships derived from average values for many individual species have slopes which are not typical for any of the component species, it is concluded that caution must be exercized when interpreting such interspecific plots in Allometric studies.     

The evolution of complex brains and behaviors in African cichlid fishes

In this review, I explore the effects of both social organization and the physical environment, specifically habitat complexity, on the brains and behavior of highly visual African cichlid fishes, drawing on examples from primates and birds where appropriate. In closely related fishes from the monophyletic Ectodinii clade of Lake Tanganyika, both forces influence cichlid brains and behavior. Considering social influences first, visual acuity differs with respect to social organization (monogamy versus polygyny). Both the telencephalon and amygdalar homologue, area Dm, are larger in monogamous species. Monogamous species are found to have more vasotocin-immunoreactive cells in the preoptic area of the brain. Habitat complexity also influences brain and behavior in these fishes. Total brain size, telencephalic and cerebellar size are positively correlated with habitat complexity. Visual acuity and spatial memory are enhanced in cichlids living in more complex environments. However habitat complexity and social forces affect cichlid brains differently. Taken together, our field data and plasticity data suggest that some of the species-specific neural effects of habitat complexity could be the consequence of the corresponding social correlates. Environmental forces, however, exert a broader effect on brain structures than social ones do, suggesting allometric expansion of the brain structures in concert with brain size and/or co-evolution of these structures [Current Zoology 56 (1): 144-156 2010].     

The maximal body mass–area relationship in island mammals

Aim A positive power relationship between maximal body mass and land area has previously been reported of the form M_max ∝ Area^0.5 whilst allometric scaling theory predicts either M_max ∝ Area^1.33 or M_max ∝ Area¹. We provide an analysis of the maximal mass–area relationship for four island systems, to test the hypothesis that community relaxation following isolation converges in each case to a slope of Area^0.5. Location Islands of the Japanese archipelago, the western Mediterranean, the Sea of Cortés and Southeast Asia. Methods We calculated the relationship between island area and the maximal body mass of the largest mammal species on the island using linear regression models with log‐transformed variables, and tested the hypothesis that the slopes were not significantly different from 0.5. Results We found a slope of 0.47 within the Japanese archipelago, 0.42 for western Mediterranean islands, 0.73 for the Sea of Cortés islands and 0.50 for Southeast Asian islands. None of these slopes were significantly different from 0.5. Main conclusions Our results provide further empirical support for previous findings of a general maximal body mass–area relationship of M_max ∝ Area^0.5, but they deviate from theoretical predictions. We hypothesize that this mass–area relationship was the ultimate end point of community relaxation initiated by the isolation of the mammal communities. Maximal body mass on each island today probably reflects the interaction between energetic constraints, home range size and island area.     

Gregarious desert locusts have substantially larger brains with altered proportions compared with the solitarious phase

The behavioural demands of group living and foraging have been implicated in both evolutionary and plastic changes in brain size. Desert locusts show extreme phenotypic plasticity, allowing brain morphology to be related to very different lifestyles in one species. At low population densities, locusts occur in a solitarious phase that avoids other locusts and is cryptic in appearance and behaviour. Crowding triggers the transformation into the highly active gregarious phase, which aggregates into dense migratory swarms. We found that the brains of gregarious locusts have very different proportions and are also 30 per cent larger overall than in solitarious locusts. To address whether brain proportions change with size through nonlinear scaling (allometry), we conducted the first comprehensive major axis regression analysis of scaling relations in an insect brain. This revealed that phase differences in brain proportions arise from a combination of allometric effects and deviations from the allometric expectation (grade shifts). In consequence, gregarious locusts had a larger midbrain∶optic lobe ratio, a larger central complex and a 50 per cent larger ratio of the olfactory primary calyx to the first olfactory neuropile. Solitarious locusts invest more in low-level sensory processing, having disproportionally larger primary visual and olfactory neuropiles, possibly to gain sensitivity. The larger brains of gregarious locusts prioritize higher integration, which may support the behavioural demands of generalist foraging and living in dense and highly mobile swarms dominated by intense intraspecific competition.     

Expression of recombinant goldfish glutamic acid decarboxylase 65 and evidence for differential pH and PLP responsiveness compared to the human enzyme

Variability of taurine (2-aminoethane sulfonic acid) was studied as a function of size in the mussel Mytilus galloprovincialis and tissue specificity. Isometric and/or allometric relationships were established with regard to total soft mass of the mussels between 20 and 60 mm shell length. Relative amounts of taurine dropped significantly with increasing mass of whole soft tissues with an allometric coefficient value of -0.15. The inverse relationship between taurine and increasing size of mussels was confirmed for gill epithelium and labial palp (allometric coefficient values of -0.16 and -0.10, respectively), tissues that, in turn, represented isometric functions with regard to total soft mass. Although relative amounts of taurine were significantly different in digestive gland, mantle and foot, relationships with increasing size of mussels remained unchanged in these tissues. Gill area of mussels was related to soft mass with an allometric coefficient of 0.70 by 2D Image Analysis, but increased to 0.85 when introducing a third dimension, i.e. gill thickness. Results are discussed according to gill structure analysis and taurine functionality.     

Causes of variation in wing loading among Drosophila species

We examined influences on wing and body size in 11 species (12 strains) of Drosophila. Six measures of wing length and width were closely correlated with wing area and suggested little variation in wing shape among the species. Among ten species wing loading, an important factor in flight costs and manoeuvrability, increased as body mass increased at a rate consistent with expectations from allometric scaling of wing area and body mass to body length. Intraspecific variation in wing loading showed similar relationships to body mass. Density and temperature during larval development influenced wing loading through general allometric relations of body size and wing area. Temperature during the pupal stage, but not during wing hardening after eclosion, influenced wing area independently of body size. Wing area increased as growth temperature decreased. Individuals reared at cooler temperatures thus compensated for a potential allometric increase in wing loading by differentially enlarging the wing area during pupal development.     

Evolution of brain–body allometry in Lake Tanganyika cichlids

Brain size is strongly associated with body size in all vertebrates. This relationship has been hypothesized to be an important constraint on adaptive brain size evolution. The essential assumption behind this idea is that static (i.e., within species) brain–body allometry has low ability to evolve. However, recent studies have reported mixed support for this view. Here, we examine brain–body static allometry in Lake Tanganyika cichlids using a phylogenetic comparative framework. We found considerable variation in the static allometric intercept, which explained the majority of variation in absolute and relative brain size. In contrast, the slope of the brain–body static allometry had relatively low variation, which explained less variation in absolute and relative brain size compared to the intercept and body size. Further examination of the tempo and mode of evolution of static allometric parameters confirmed these observations. Moreover, the estimated evolutionary parameters indicate that the limited observed variation in the static allometric slope could be a result of strong stabilizing selection. Overall, our findings suggest that the brain–body static allometric slope may represent an evolutionary constraint in Lake Tanganyika cichlids.     

Seed beetle survivorship,growth and egg size plasticity in a paloverde hybrid zone

1. Although numerous studies have examined the ecology of plant resistance to herbivores and the distribution of herbivores within plant hybrid zones, few have examined how plant hybridization influences herbivore growth, development, or life history. The experiment reported here examines variation in survivorship, development time, and final adult body size of Stator limbatus reared on seeds of parental and hybrid Cercidium floridum-×-C. microphyllum trees from a paloverde hybrid zone in eastern California. Because S. limbatus exhibits egg size plasticity in response to host species, the size of eggs that females lay on hybrid and parental plants is also examined. 2. The hypotheses (a) that seeds of hybrid trees are intermediate between those of the two parental species in their resistance to penetration by S. limbatus larvae; (b) that seeds of hybrid trees are intermediate in their suitability for the growth of larvae that successfully penetrate the seed-coat; (c) that female S. limbatus can distinguish between hybrid trees and pure-bred trees, as quantified by the size of eggs they lay on seeds of each taxon, and (d) that female S. limbatus can distinguish among individual hybrid trees, are tested. 3. On average, S. limbatus survivorship was lower, development time longer, and emergence body mass lower on seeds of C. floridum than on seeds of C. microphyllum. Seeds of hybrid trees were, on average (across trees), intermediate between seeds of the parental species in the resistance of their seed-coats to penetration by S. limbatus larvae and in their suitability for larval growth. Individual hybrid trees also varied in the resistance of their seeds to, and the suitability of their seeds for, S. limbatus larvae. 4. Female S. limbatus laid significantly larger eggs on seeds of C. floridum than on seeds of C. microphyllum, and, on average, intermediate size eggs on hybrid trees. The size of eggs laid by females also varied among hybrid trees, with females laying C. floridum-sized eggs on some trees, and C. microphyllum-sized eggs on other trees. These results suggest that females have at least some ability to distinguish among hybrid trees, but there was no evidence that females laid larger eggs on poorer quality hybrid trees.     

Variability and size in mammals and birds

Body size, its variability, and their ecological correlates have long been important topics in evolutionary biology. Yet, the question of whether there is a general relationship between size and size-relative variability has not previously been addressed. Through an analysis of body-mass and length measurements from 65 074 individuals from 351 mammalian species, we show that size-relative variability increases significantly with mean species body size. Analysis of mean body mass and standard deviations for 237 species of birds revealed the same pattern. We present three plausible alternatives explanations and eliminate several others. Of these, the hypothesis that the increase in size-relative variability with mean body mass is related to the scaling of body mass components is most strongly supported. In effect, larger mammals and birds are more variable because their body mass is composed to greater relative degree of components with higher intrinsic variability (bone, fat, and muscle). In contrast, smaller mammals and birds have lower body mass variability because they are composed to a greater relative extent of components (viscera and nervous system) in which size variation is more highly constrained by energetic and functional factors.     

Patterns of sexual size dimorphism in cattle breeds support Rensch’s rule

Rensch’s rule describes the pattern of sexual size dimorphism (SSD) claiming that in taxa where males are the larger sex, larger species generally exhibit higher male to female body size ratios. Agreement with Rensch’s rule is manifested by the slope of the allometric relationship between male and female body size exceeding one. In this paper we have tested the hypothesis that recent rapid evolutionary changes of body size accompanying domestication process and morphological radiation of domestic breeds follow Rensch’s rule. We have analyzed literature data on adult body size of males and females in domestic cows, yaks, buffaloes and other bovines (315, 12, 24 and 2 breeds, respectively) and compared it with SSD in 18 wild species/subspecies of the subfamily Bovinae. Male to female body mass ratio in domestic cows (1.48) was fairly comparable to that found in other species of domestic and wild bovines except domestic buffaloes (1.19). In cows we have demonstrated clear positive allometry of male to female body mass ratio (slope 1.21) predicted by Rensch’s rule, however, no such clear relationship was found when body mass was replaced by shoulder height. These findings are in agreement with those we have previously reported in other livestock species, goats and sheep.