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opencc-zeroDec 2016View details →

Data from: Early postnatal exposure to airborne fine particulate matter induces autism-like phenotypes in male rats

Epidemiological studies have revealed that ambient fine particulate matter (PM2.5) exposure is closely associated with autism spectrum disorder (ASD). However, there is a relative paucity of laboratory data to support this epidemic finding. In order to assess the relationship between PM2.5 exposure and ASD, neonatal male Sprague-Dawley (SD) rats were chosen and exposed to PM2.5 (2 or 20 mg/kg body weight, once a day) by intranasal instillation from postnatal day (PND) 8 to 22. It was found that when exposed to PM2.5 in the early neonatal period for two weeks, both groups of the exposure rats manifested typical behavioral features of autism, including communication deficits, poor social interaction and novelty avoidance. And, we further found, among five ASD candidate genes we chose, both the mRNA level and protein expression of SH3 and multiple ankyrin repeat domains 3 (Shank3) decreased significantly in the rat hippocampus after high dose of PM2.5 exposure. Moreover, results showed that PM2.5-exposure significantly increased the levels of pro-inflammatory cytokines, IL-1β, IL-6, and TNF-α in the hippocampus and prefrontal cortex. The expression of Glial fibrillary acidic protein (GFAP) and ionized binding calcium adapter molecule (IBA1), markers of astrocytes and microglial cell activation, respectively, also increased in the exposed animals. Our work provides new data on the link between postnatal exposure to ambient PM2.5 and the onset of ASD-like symptoms in human beings, and the increased inflammatory response and abnormalities in Shank3 expression in the brain may contribute to the mechanisms of PM2.5 exposure induced ASD.

opencc-zeroDec 2014View details →

Data from: Using the Phenoscape Knowledgebase to relate genetic perturbations to phenotypic evolution

The abundance of phenotypic diversity among species can enrich our knowledge of development and genetics beyond the limits of variation that can be observed in model organisms. The Phenoscape Knowledgebase (KB) is designed to enable exploration and discovery of phenotypic variation among species. Because phenotypes in the KB are annotated using standard ontologies, evolutionary phenotypes can be compared with phenotypes from genetic perturbations in model organisms. To illustrate the power of this approach, we review the use of the KB to find taxa showing evolutionary variation similar to that of a query gene. Matches are made between the full set of phenotypes described for a gene and an evolutionary profile, the latter of which is defined as the set of phenotypes that are variable among the daughters of any node on the taxonomic tree. Phenoscape's semantic similarity interface allows the user to assess the statistical significance of each match and flags matches that may only result from differences in annotation coverage between genetic and evolutionary studies. Tools such as this will help meet the challenge of relating the growing volume of genetic knowledge in model organisms to the diversity of phenotypes in nature. The Phenoscape KB is available at http://kb.phenoscape.org.

opencc-zeroDec 2008View details →

Data from: The genetics of phenotypic plasticity in plant defense: trichome production in Mimulus guttatus

Insect herbivory is a major driving force of plant evolution. Phenotypic plasticity and developmental variation provide a means for plants to cope with variable herbivory. We characterized the genetics of developmental variation and phenotypic plasticity in trichome density, a putative defensive trait of Mimulus guttatus (yellow monkeyflower). Our results are evaluated in relation to the optimal defense theory, which provides testable predictions for plastic and developmental patterns in defense traits. We found that both developmental stage and simulated insect damage affected trichome production, but in different ways. Plants were more likely to produce at least some trichomes on later leaves than on earlier leaves, regardless of damage. Damage did not affect the average probability of producing trichomes, but it did increase the density of hairs on trichome‐positive plants. We mapped trichome quantitative trait loci (QTL) by selectively genotyping a large panel of recombinant inbred lines derived from two highly divergent populations. Several highly pleiotropic QTL influenced multiple aspects of the trichome phenotype (constitutive, developmental, and/or plastic responses). Only one of the QTL influenced trichome induction following damage. In a result that is consistent with a central prediction of optimal defense theory, the high allele at this location was from the ancestral population with low constitutive trichome production.

Data from: Phenotypic biomarkers of climatic impacts on declining insect populations: a key role for decadal drought, thermal buffering and amplification effects and host plant dynamics

1. Widespread population declines have been reported for diverse Mediterranean butterflies over the last three decades, and have been significantly associated to increased global change impacts. The specific landscape and climatic drivers of these declines remain uncertain for most declining species. 2. Here we analyse whether plastic phenotypic traits of a model butterfly species (Pieris napi) perform as reliable biomarkers of vulnerability to extreme temperature impacts in natural populations, showing contrasting trends in thermally exposed and thermally buffered populations. 3. We also examine whether improved descriptions of thermal exposure of insect populations can be achieved by combining multiple information sources (i.e. integrating measurements of habitat thermal buffering, habitat thermal amplification, host plant transpiration, and experimental assessments of thermal death time (TDT), thermal avoidance behaviour (TAB) and thermally induced trait plasticity). These integrative analyses are conducted in two demographically declining and two non-declining populations of P. napi. 4. The results show that plastic phenotypic traits (butterfly body mass and wing size) are reliable biomarkers of population vulnerability to extreme thermal conditions. Butterfly wing size is strongly reduced only in thermally exposed populations during summer drought periods. Lab rearing of these populations documented reduced wing size due to significant negative effects of increased temperatures affecting larval growth. We conclude that these thermal biomarkers are indicative of the population vulnerability to increasing global warming impacts, showing contrasting trends in thermally exposed and buffered populations. 5. Thermal effects in host plant microsites significantly differ between populations, with stressful thermal conditions only effectively ameliorated in mid-elevation populations. In lowland populations we observe a six-fold reduction in vegetation thermal buffering effects, and larval growth occurs in these populations at significantly higher temperatures. Lowland populations show reduced host plant quality (C/N ratio), reduced leaf transpiration rates and complete aboveground plant senescence during the peak of summer drought. Amplified host plant temperatures are observed in open microsites, reaching thermal thresholds that can affect larval survival. 6. Overall, our results suggest that butterfly population vulnerability to long-term drought periods is associated to multiple co-occurring and interrelated ecological factors, including limited vegetation thermal buffering effects at lowland sites, significant drought impacts on host plant transpiration and amplified leaf surface temperature, as well as reduced leaf quality linked to the seasonal advance of plant phenology. Our results also identify multi-annual summer droughts affecting larval growing periods as a key driver of the recently reported butterfly population declines in the Mediterranean biome.

Data from: Many-to-many mapping of phenotype to performance: an extension of the F-matrix for studying functional complexity

Performance capacity influences ecology, behavior and fitness, and is determined by the underlying phenotype. The phenotype-performance relationship can influence the evolutionary trajectory of an organism. Several types of phenotype-performance relationships have been described, including one-to-one relationships between a single phenotypic trait and performance measure, trade-offs and facilitations between a phenotypic trait and multiple performance measures, and redundancies between multiple phenotypic traits and a single performance measure. The F-matrix is an intraspecific matrix of measures of statistical association between phenotype and performance that is used to quantify these relationships. We extend the F-matrix in two ways. First, we use the F-matrix to describe how the different phenotype-performance relationships occur simultaneously and interact in functional systems, a phenomenon we call many-to-many mapping. Second, we develop methods to compare F-matrices among species and compare phenotype-performance relationships at microevolutionary and macroevolutionary levels. We demonstrate the expanded F-matrix approach with a dataset of eight phrynosomatine lizard species, including six phenotypic traits and two measures of locomotor performance. Our results suggest that all types of relationships occur in this system and that phenotypic traits involved in trade-offs are more functionally constrained and tend evolve slower interspecifically than those involved in facilitations or one-to-one relationships.

Data from: "Alternative phenotypes within mating systems" in The evolution of insect mating systems

Insects were for a long time considered simple organisms with unvarying behavioural repertoires, incapable of complicated behavioural responses to changing environments and/or social conditions. However, nothing could be further from the truth; phenotypic plasticity is widespread in insect development, life history, physiology, and behaviour (Whitman and Ananthakrishnan 2009). Plastic responses to environmental and social conditions are actually central to the remarkable adaptability of insects, and have played a crucial role in their evolutionary histories (Moczek 2010; Simpson et al. 2011). Moreover, phenotypic plasticity in insects is not merely restricted to simple responses in metabolism or activity to abiotic factors such as temperature, but can be extremely elaborate, an illuminating example of which is the learning ability of honeybees (Giurfa 2007; Hammer and Menzel 1995; Menzel 1993; Menzel and Muller 1996). Insect mating systems are no exception to this pattern. This chapter explores the intrasexual variation in behaviours and morphologies found in insect mating systems. More specifically, we focus on the evolution of the alternative means by which individuals obtain fertilizations, generally referred to as 'alternative mating tactics', or 'alternative mating phenotypes' (AMPs). The first studies to describe what we can interpret today as cases of AMPs in insects date back to at least the 1930s (Salt 1937), but it was only in the 1970s that the number of studies reporting this phenomenon started to accumulate. In 1983, when the classic examples of AMPs in digger bees and scorpionflies were reviewed by Thornhill and Alcock (1983), approximately 50 cases of AMPs in insects were already known, a number that has now surpassed the 200 mark. Here, we review the theoretical and empirical advances that have been made in this area since Thornhill and Alcock's volume. We start by describing two illustrative systems in detail, gryllid field crickets and onthophagine dung beetles. These two groups were chosen because their reproductive biology is well known, and because the contrasting degrees of behavioural plasticity and morphological specialisation between alternative phenotypes in these two groups illustrate the diversity of AMPs that has evolved in insects. We then discuss the genetic models that have been proposed to account for the evolution and maintenance of such dimorphisms, before reviewing the occurrence of AMPs in insects more generally. Finally, we discuss the relatively limited evidence for AMPs in female insects, a somewhat new and very promising area for future research.

Data from: Sweet tetra-trophic interactions: multiple evolution of nectar secretion, a defensive extended phenotype in cynipid gall wasps

Many herbivores employ reward-based mutualisms with ants to gain protection from natural enemies. We examine the evolutionary dynamics of a tetra-trophic interaction in which gall wasp herbivores induce their host oaks to produce nectar-secreting galls, which attract ants that provide protection from parasitoids. We show that, consistent with other gall defensive traits, nectar secretion has evolved repeatedly across the oak gall wasp tribe and also within a single genus (Disholcaspis) that includes many nectar-inducing species. Once evolved, nectar secretion is never lost in Disholcaspis, consistent with high defensive value of this trait. We also show that evolution of nectar secretion is correlated with a transition from solitary to aggregated oviposition, resulting in clustered nectar-secreting galls, which produce a resource that ants can more easily monopolize. Such clustering is commonly seen in ant guard mutualisms. We suggest that correlated evolution between maternal oviposition and larval nectar induction traits has enhanced the effectiveness of this gall defense strategy.

Data from: Maternal effects influence phenotypes and survival during early life stages in an aquatic turtle

Offspring phenotypic variation can be substantially influenced by non-genetic factors such as maternal effects, which ultimately can influence organismal fitness. For oviparous organisms that lack parental care, oviposition-site choice and egg size are maternal effects that can greatly affect offspring traits. Yet, few studies examine the consequences of these traits in the wild. We manipulated the contents of natural painted turtle nests such that offspring spent two life stages (incubation and hibernation) in either maternally-selected nest sites or randomly-selected nest sites, and quantified treatment differences in environmental parameters and offspring phenotypes. Additionally, we tracked the fates of individual eggs and hatchlings, which allowed us to quantify the strength and form of selection acting on egg size during incubation and, for the first time, body size during hibernation. Maternally-selected nest sites were warmer and produced offspring that were longer and hatched earlier than their siblings emerging from cooler, randomly-selected nests. Treatments did not affect any measured traits during hibernation. We detected no selection on egg size during the incubation stage, but significant linear selection favoring larger hatchlings during hibernation. Our results suggest that nest-site choice allows mothers to partially control the environment of their incubating eggs, but is less effective at controlling hatchling environments during hibernation. Additionally, we provide novel support for the "bigger-is-better" hypothesis in turtles by showing a positive relationship between size and survival during the hibernation stage.

Data from: Are thyroid hormones mediators of incubation temperature-induced phenotypes in birds?

Incubation temperature influences a suite of traits in avian offspring. However, the mechanisms underlying expression of these phenotypes are unknown. Given the importance of thyroid hormones in orchestrating developmental processes, we hypothesized that they may act as an upstream mechanism mediating the effects of temperature on hatchling phenotypic traits such as reduced growth and thermoregulation. We found that plasma T3, but not T4 concentrations, differed among newly-hatched wood ducks (Aix sponsa) from different embryonic incubation temperatures. T4 at hatching correlated with time spent hatching, and T3 correlated with hatchling body condition, tarsus length, time spent hatching, and incubation period. In addition, the T3:T4 ratio differed among incubation temperatures up at hatch. Our findings are consistent with the hypothesis that incubation temperature modulates plasma thyroid hormones which in turn influences multiple aspects of duckling phenotype.

opencc-zeroDec 2016View details →

Data from: Adaptive shaping of the behavioural and neuroendocrine phenotype during adolescence

Environmental conditions during early life can adaptively shape the phenotype for the prevailing environment. Recently, it has been suggested that adolescence represents an additional temporal window for adaptive developmental plasticity, though supporting evidence is scarce. Previous work has shown that male guinea pigs living in large mixed-sex colonies develop a low-aggressive phenotype as part of a queuing strategy that is adaptive for integrating into large unfamiliar colonies. By contrast, males living in pairs during adolescence become highly aggressive towards strangers. Here, we tested whether the high-aggressive phenotype is adaptive under conditions of low population density, namely when directly competing with a single opponent for access to females. For that purpose, we established groups of one pair-housed male (PM), one colony-housed male (CM) and two females. PMs directed more aggression towards the male competitor and more courtship and mating towards females than did CMs. In consequence, PMs attained the dominant position in most cases and sired significantly more offspring. Moreover, they showed distinctly higher testosterone concentrations and elevated cortisol levels, which probably promoted enhanced aggressiveness while mobilizing necessary energy. Taken together, our results provide the clearest evidence to date for adaptive shaping of the phenotype by environmental influences during adolescence.

Data from: Functional traits in red flour beetles: the dispersal phenotype is associated with leg length but not body size nor metabolic rate

Individuals vary in their ability to disperse. Much of this variation can be described by covarying phenotypic traits that are related to dispersal (constituting the 'dispersal phenotype' or 'dispersal syndrome'), but the nature of the associations among these traits is not well understood. Unravelling the associations among traits that potentially constitute the dispersal phenotype provides a foundation for understanding evolutionary trade-offs due to variation in dispersal. Here, we tested five predictions pertaining to the relationships among physiological, morphological and movement traits that are associated with dispersal, using a species with a long history as a laboratory model for studying ecological phenomena, red flour beetles (Tribolium castaneum). We identified a dominant axis of movement ability that describes variation in dispersal-related movement traits. Individuals that scored positively on this axis moved at higher speed, travelled longer distances, had lower movement intermittency and dispersed quicker to a specified area. Relative leg length, but not body size nor routine metabolic rate related positively with movement ability, indicating a likely mechanistic relationship between increased stride length and movement ability. Our data suggest that the dispersal phenotype may be more strongly linked to morphological traits than physiological ones. We demonstrate that associations among many functional traits do not necessarily conform to a priori expectations, and predict that the substantial intraspecific variation in trait values may be important for selection. Movement is a complex behavioural trait, but it has a mechanistic basis in locomotor morphology that warrants further exploration.

Data from: Theoretical conditions for the coexistence of viral strains with differences in phenotypic traits: a bifurcation analysis

We investigate the dynamics of a wild-type viral strain which generates mutant strains differing in phenotypic properties for infectivity, virulence, and mutation rates. We study, by means of a mathematical model and bifurcation analysis, conditions under which the wild-type and mutant viruses, which compete for the same host cells, can coexist. The coexistence conditions are formulated in terms of the basic reproductive numbers of the strains, a maximum value of the mutation rate, and the virulence of the pathogens. The analysis reveals that parameter space can be divided into five regions, each with distinct dynamics, that are organised around degenerate Bogdanov-Takens and zero-Hopf bifurcations, the latter of which gives rise to a curve of transcritical bifurcations of periodic orbits. These results provide new insights into the conditions by which viral populations may contain multiple coexisting strains in a stable manner.

Data from: Experimental evolution reveals balancing selection underlying coexistence of alternative male reproductive phenotypes

Heritable alternative reproductive phenotypes (ARPs), which differ in traits associated with competition for mates, occur across taxa. If polymorphism in the genes underlying ARPs is maintained by balancing selection, selection should return ARP proportions to their equilibrium if that equilibrium is perturbed. Here, we used an experimental evolution approach to directly test this prediction in male Rhizoglyphus robini, in which two heritable morphs occur: armoured fighters and more female-like, benign scramblers. Using selection lines nearly fixed for male morph, we constructed replicate populations consisting of 50% or 94% fighters, and allowed them to evolve for 14 generations in two types of environment: simple or spatially complex. We found that in both types of populations, the proportion of fighters converged on values within a narrow range of 0.70-0.83, although the rate of convergence was slower in the complex environment. Our results thus demonstrate balancing selection acting on polymorphism(s) underlying ARPs

Data from: There's no place like home: the contribution of direct and extended phenotypes on the expression of spider aggressiveness

Architectural constructions allow animals to modify their environment in order to improve their reproductive success. Constructions also modulate the expression of individual behavior, ultimately affecting the presence and importance of animal personality within populations. The exact impact of constructions on personality is seldom investigated. We quantified experimentally the impact of web characteristics on individual foraging behavior in the Western black widow spider (Latrodectus hesperus). We assayed aggressiveness toward a prey cue, and boldness while individuals resided on their own web versus after being translocated onto webs built by con-specifics. We quantified the importance of individual differences in aggressiveness and boldness while accounting for differences in web characteristics. We tested for relationships between web building, aggressiveness, and boldness. Web characteristics affected spider aggressiveness and interacted with individual web-building behavior to explain up to a fifth of the variation in foraging aggression, but did not affect spider boldness. Even after accounting for web characteristics, individuals still exhibited important differences in aggressiveness. We detected no relationship between an individual's aggressiveness behavioral type and the characteristics of the web it built. Surprisingly, web characteristics impacted aggressiveness differently from one individual to the next. Hence the effect of web characteristics on foraging behavior might depend on condition or past experience. Webs contributed mostly to the variation in aggressiveness within-individuals. Variation in web building behavior might affect the amount of consistent differences in foraging, mating, and anti-predator behavior among individuals and needs to be accounted for when quantifying individual variation in behavior in spiders.

opencc-zeroMay 2019View details →

Data from: Introduced parasite changes host phenotype, mating signal and hybridisation risk: Philornis downsi effects on Darwin's finch song

Introduced parasites that alter their host's mating signal can change the evolutionary trajectory of a species through sexual selection. Darwin's Camarhynchus finches are threatened by the introduced fly Philornis downsi that is thought to have accidentally arrived on the Galapagos Islands during the 1960s. The P. downsi larvae feed on the blood and tissue of developing finches causing on average ~55 % in-nest mortality and enlarged naris size in survivors. Here we test if enlarged naris size is associated with song characteristics and vocal deviation in the small tree finch (Camarhynchus parvulus), the critically endangered medium tree finch (C. pauper), and the recently observed hybrid tree finch group (Camarhynchus hybrids). Male C. parvulus and C. pauper with enlarged naris size produced song with lower maximum frequency and greater vocal deviation, but there was no significant association in hybrids. Less vocal deviation predicted faster pairing success in both parental species. Finally, C. pauper males with normal naris size produced species-specific song, but male C. pauper with enlarged naris size had song that was indistinguishable from other tree finches. When parasites disrupt host mating signal, they may also facilitate hybridisation. Here we show how parasite-induced naris enlargement affects vocal quality resulting in blurred species mating signals.

Data from: Hatching hierarchy but not egg-related effects governs behavioral phenotypes in gull chicks

In many bird species that practice parental care, siblings often compete for resources and care provided by their parents, although their strategies differ according to hatching rank and condition. Differences in offspring strategies are generally attributed to hatching order and maternal effects, which are difficult to separate because these effects are often correlated. For example, third-hatched chicks of large gull species receive more egg testosterones and corticosterone, which influence early behavioral patterns. In this study, we carried out a cross-fostering experiment with first- and last-laid eggs of the yellow-legged gull (Larus michahellis) to test whether the within-brood variation in behavioral strategies for competing with siblings and coping with stress are due to maternal effects or to hatching order. Chicks hatched in the last position within the experimental brood emitted more chatter calls to attract parents' attention, were less prone to respond to warning of danger, and had a lower breathing rate while restrained than first-hatched chicks. Egg laying order did not affect chick behaviors or breathing rate. Thus, we concluded that the different behavioral strategies of chicks were determined by their posthatching experience and not by the original egg position within the clutch. Last-laid eggs were smaller and chicks from those eggs grew slower than chicks from first-laid eggs. Independently of the original laying order, chicks that hatched first in the experimental brood grew faster than their siblings. Overall, our results indicate that behavioral strategies of chicks are plastic and influenced by their early social experience.

Data from: A costly chemical trait: phenotypic condition dependence of cuticular hydrocarbons in a dung beetle

Chemical traits are increasingly recognised as important cues used in mate choice. For example, the cuticular hydrocarbons (CHCs) of insects have been shown to influence mating success in a range of taxa. Less is known, however, about how CHCs are expressed in proportion to an individual's condition, and consequently whether CHCs can function as condition dependent signals of quality. We investigated this question using the dung beetle, Onthophagus taurus. CHCs are subject to sexual selection in this species through mate choice. A dietary manipulation revealed condition dependence of CHC expression for both sexes: dietary restriction decreased overall CHC production and altered the composition of CHCs. Furthermore, CHC production was associated with a measure of condition in beetles fed a limited diet but not those fed ad libitum. These results implicate a resource cost to CHC production that is likely to result in trade-offs with other fitness components in this species, as these respond similarly to a dietary restriction. The CHC profiles showed sexual dimorphism: males produced more CHCs and the sexes differed in the blend of compounds they produced. There was evidence for a male dimorphism in the CHC profile, in line with the presence of the alternative reproductive tactics (minor sneaks and major fighters) in this species. However, rather than mimicking a female CHC profile, minor males differed more from females than major males. Our results suggest that CHCs are a costly trait in O. taurus that has the potential to act as a condition dependent signal of quality.

opencc-zeroJun 2021View details →

Variability, heritability and condition-dependence of the multidimensional male colour phenotype in a passerine bird

Elaborate ornamental traits are commonly assumed to be honest signals of individual quality, owing to the presumed costs involved in their production and/or maintenance. Such traits are often highly variable, possibly because of condition-dependence and/or high underlying genetic variation, and it has been suggested that their expression should be more sensitive to condition and/or more heritable than non-ornamental traits. Many bird species display colourful plumage with multiple distinct patches of different developmental origins, forming complex colour phenotypes. Despite this complexity, colourful ornaments are often studied in isolation, without comparison to suitable non-ornamental controls. Based on plumage reflectance data collected over 8 years, we assessed the signalling potential of the multidimensional male colour phenotype in a tropical bird: the purple-crowned fairy-wren Malurus coronatus. Specifically, we tested the predictions that the express ion of putative ornamental colours (purple and black –the breeding colours– and blue) is (1) more variable, (2) more heritable and (3) more condition-dependent compared to year-round non-ornamental colours (buff-white and brown). Our results show that ornamental colours exhibit greater levels of variability, and some chromatic components of purple and blue colouration appear slightly heritable (h² = 0.19-0.30). However, contrary to predictions of heightened condition-dependence in ornaments, only brightness of the buff-white and brown colouration increased with male body condition, although brightness of the purple colouration was related to male age as expected. Despite partial support for predictions, the lack of consistent patterns illustrates the complexity of visual signals and highlights the need to study colour phenotypes in their entirety.

Data from: Thrifty phenotype vs cold adaptation: trade-offs in upper limb proportions of Himalayan populations of Nepal

The multi-stress environment of high altitude has been associated with growth deficits in humans, particularly in zeugopod elements (forearm, lower leg). This is consistent with the thrifty phenotype hypothesis, which has been observed in Andeans, but has yet to be tested in other high altitude populations. In Himalayan populations, other factors, such as cold stress, may shape limb proportions. The current study investigated whether relative upper limb proportions of Himalayan adults (n=254) differ between highland and lowland populations, and whether cold adaptation or a thrifty phenotype mechanism may be acting here. Height, weight, humerus length, ulna length, hand length, and hand width were measured using standard methods. Relative to height, total upper limb and ulna lengths were significantly shorter in highlanders compared to lowlanders in both sexes, whilst hand and humerus length were not. Hand width did not significantly differ between populations. These results support the thrifty phenotype hypothesis, as hand and humerus proportions are conserved at the expense of the ulna. The reduction in relative ulna length could be attributed to cold adaptation, but the lack of difference between populations in both hand length and width indicate that cold adaptation is not shaping hands proportions in this case.