Sex-specific developmental plasticity in response to yolk corticosterone in an oviparous lizard

Corticosterone exposure during prenatal development as a result of maternal upregulation of circulating hormone levels has been shown to have effects on offspring development in mammals. Corticosterone has also been documented in egg yolk in oviparous vertebrates, but the extent to which this influences phenotypic development is less studied. We show that maternal corticosterone is transferred to egg yolk in an oviparous lizard (the mallee dragon, Ctenophorus fordi Storr), with significant variation among clutches in hormone levels. Experimental elevation of yolk corticosterone did not affect hatching success, incubation period or offspring sex ratio. However, corticosterone did have a sex-specific effect on skeletal growth during embryonic development. Male embryos exposed to relatively high levels of corticosterone were smaller on average than control males at hatching whereas females from hormone-treated eggs were larger on average than control females. The data thus suggest that males are not just more sensitive to the detrimental effects of corticosterone but rather that the sexes may have opposite responses to corticosterone during development. Positive selection on body size at hatching for both sexes in this species further suggests that increased corticosterone in egg yolk may have sex-specific fitness consequences, with potential implications for sex allocation and the evolution of hormone-mediated maternal effects.

Consequences of maternal yolk testosterone for offspring development and survival : experimental test in a lizard

1. Hormone-mediated maternal effects and developmental plasticity are important sources of phenotypic variation, with potential consequences for trait evolution. Yet our understanding of the importance of maternal hormones for offspring fitness in natural populations is very limited, particularly in non-avian species.

2. We experimentally elevated yolk testosterone by injection of a physiological dose into eggs of the lizard Ctenophorus fordi Storr, to investigate its roles in offspring development, growth and survival.

3. Yolk testosterone did not influence incubation period, basic hatchling morphology or survival under natural conditions. However, there was evidence for increased growth in hatchlings from testosterone-treated eggs, suggesting that maternal hormones have potential fitness consequences in natural populations.

4. The positive effect of prenatal testosterone exposure on postnatal growth could represent a taxonomically widespread developmental mechanism that has evolved into an adaptive maternal effect in some taxa, but remains deleterious or selectively neutral in others.

5. A broader taxonomic perspective should increase our understanding of the role of physiological constraints in the evolution of endocrine maternal effects.

Responses to flooding and drying in seedlings of a common Australian desert floodplain shrub : Muehlenbeckia florulenta Meisn. (tangled lignum)

We investigated the effects of flooding and drying over 6 months on growth and biomass allocation in seedlings of Muehlenbeckia florulenta Meisn. (tangled lignum), a common and widely distributed shrub of Australia's desert floodplains. We sought to determine if lignum seedlings respond to flooding or drying by altering traits or allocation patterns or instead display fixed patterns of development. Since desert floodplains are highly unpredictable and heterogeneous environments, we hypothesised that adaptive phenotypic plasticity is unlikely to have developed or be advantageous in seedlings of this species as environmental state changes are highly variable in their timing and duration and plants risk being caught out of kilter with environmental conditions. To test this, we conducted a glasshouse experiment in which lignum seedlings, grown in both clay and sandy sediments, were subjected to a range of hydrological conditions over a period of 6 months. Lignum seedlings exhibited considerable tolerance of both flooding and drying in our experiment and no mortality was observed. Growth was significantly reduced by flooding, however, and seedlings displayed extremely delayed development rather than plasticity in overall biomass allocation or any of the specific morphological variables we measured. Lignum seedlings were considerably more tolerant of drying than flooding and responded plastically by reducing leaf area ratios through reductions in specific leaf areas and leaf production and expansion. Sediment type had little effect on seedling development. Our results indicate that surface water hydrology is likely to be a major determinant of recruitment patterns in this ecologically significant species.

Ecology of Pittosporum undulatum Vent. (Pittosporaceae) an environmental weed in south east Australia

Pittosporum undulatum Vent. (Sweet Pittosporum) is a densely foliaged tall shrub or small tree, native to the wet forests of south east Australia, This species now functions as a serious environmental weed in a range of habitats in Australia and on other continents and islands throughout the temperate, sub-tropical and tropical zones. This study investigated some of the ecological causes and consequences of P. undulatum invasion across a range of habitat types in south east Australia. Key aspects of P. undulatum biology and ecology investigated in the current study include; patterns of morphological variation across the range of habitats occupied (as a measure of the species’ plasticity), dispersal ecology and seed germinability, population structure and spatial pattern, community relationships and the ecological impacts of invasion. Phenotypic plasticity is considerable in P. undulatum. No clear patterns of geographic variation emerged from a study of leaf morphological attributes across the current range of this species on mainland south east Australia. The pattern of morphological variation is particularly complex in Victoria, where the invasion of this species is most advanced. The species’ adaptability to a range of environments and environmental conditions will likely promote further range expansion. The abundant winter fruit crop produced by functionally female P. undulatum plants attracts a suite of generalist opportunistic frugivores, which feed on P. undulatum fruits and seeds at various stages of fruit dehiscence, thereby enhancing dispersal opportunities for this species. P. undulatum seed collected from natural and invasive populations, at two stages of fruit maturity and from the scats and pellets of dispersal agents, displayed high germinability. European Blackbirds and Pied Currawongs are implicated as the main avian dispersal agents of P undulatum in south east Australia. The broader ecological implications of developing relationships between invasive fleshy-fruited bird-dispersed plant species and adaptive frugivores are likely to be considerable. The distribution of P. undulatutn seedlings was significantly negatively correlated with adult conspecifics and significantly positively correlated with trees and shrubs of other genera. This pattern reflects the importance of both firugivorous dispersal agents and the species’ germination and establishment requirements, in shaping the contagious distribution pattern typical of this species. These analyses suggest that recruitment opportunities for conspecific seedlings are limited beneath the canopy of adult conspecifics. Densities of P. undulatum were on average, 2.7 times higher in invaded populations, compared to the natural populations sampled. A male-bias was evident in all populations and no relationships between reproductive activity and the density of seedlings and juveniles were evident. Invading populations of P. undulatum impose substantial changes on ecosystem-level properties and functions. Mean species richness and cover-abundance declined notably once P. undulatum cover-abundance exceeded 20% at the invaded sites and 60% at the natural sites sampled. The natural communities sampled displayed comparatively greater resilience to the competitive effects of P. undulatum, but community attributes were affected at high densities and cover-abundance of this species. The cover-abundance of herbs and grasses declined most substantially with increasing P. undulatum at invaded sites, whereas, at the natural sites sampled, the species’ structural analogues appeared to be most affected by increasing P. undulatum cover-abundance. This study has demonstrated that the ecological consequences of P. undulatum population expansion are substantial and contribute to changes in the composition and successional trajectory of affected communities. These processes ultimately lead to the loss and simplification of biodiversity values and the homogenisation of affected habitats. P. undulatum has the potential to emerge as one of south east Australia's most serious environmental weed species.

Burial depth distribution of fennel pondweed tubers (Potamogeton pectinatus) in relation to foraging by Bewick's swans

Deep burial in the sediment of tubers of fennel pondweed (Potamogeton pectinatus) has been explained in terms of avoidance by escape against consumption by Bewick's swans (Cygnus columbianus bewickii) in autumn. We therefore expected changes in foraging pressure to ultimately result in a change in the tuber distribution across sediment depth. A trade-off underlies this idea: deep tubers are less accessible to swans but must be larger to meet the higher energy demands of sprouting in spring. To test this prediction, we compared tuber burial depth over a gradient of foraging pressure both across space and across time. Tuber samples were obtained after aboveground plant senescence but before arrival of Bewick's swans. First, we compared the current tuber bank depth profile in a shallow lake with high foraging pressure, the Lauwersmeer, with that in two wetlands with moderate and low foraging pressure. Second, we compared the current tuber burial in the Lauwersmeer with that in the early 1980s when exploitation by swans had just started there. In accordance with our hypothesis, we found significantly deeper burial of tubers under high consumption risk compared to low consumption risk, both when comparing sites and comparing time periods. Since tubers in effect only survive to the next spring, the observed differences in burial depth among sites and over time cannot be a direct result of tuber losses due to consumption by swans. Rather, these observations suggest adaptive responses in tuber burial related to foraging pressure from Bewick's swans in the recent past. We thus propose that fennel pondweed exhibits flexible avoidance by escape, of a kind rarely described for plants, where both phenotypic plasticity and genotype sorting may contribute to the observed differences in tuber burial.

Aquatic plant shows flexible avoidance by escape from tuber predation by swans

Deeper burial of bulbs and tubers has been suggested as an escape against below-ground herbivory by vertebrates, but experimental evidence is lacking. As deep propagule burial can incur high costs of emergence after dormancy, burial depth may represent a trade-off between sprouting survival and herbivore avoidance. We tested whether burial depth of subterraneous tubers is a flexible trait in fennel pondweed (Potamogeton pectinatus), facing tuber predation by Bewick's swans (Cygnus columbianus bewickii) in shallow lakes in winter. In a four-year experiment involving eight exclosures, winter herbivory by swans and all vertebrate summer herbivory were excluded in a full-factorial design; we hence controlled for aboveground vertebrate herbivory in summer, possibly influencing tuber depth. Tuber depth was measured each September before swan arrival and each March before tuber sprouting. In accordance with our hypothesis, tuber depth in September decreased after excluding Bewick's swans in comparison to control plots. The summer exclosure showed an increase in tuber biomass and the number of shallow tubers, but not a significant effect on the mean burial depth of tuber mass. Our results suggest that a clonal plant like P. pectinatus can tune the tuber burial depth to predation pressure, either by phenotypic plasticity or genotype sorting, hence exhibiting flexible avoidance by escape. We suggest that a flexible propagule burial depth can be an effective herbivore avoidance strategy, which might be more widespread among tuber forming plant species than previously thought.

Introgression of domesticated alleles into a wild trout genotype and the impact on seasonal survival in natural lakes

We tested the fitness consequences of introgression of fast-growing domesticated fish into a wild population. Fry from wild and domesticated rainbow trout (Oncorhynchus mykiss) crosses, F1 hybrids, and first- and second-generation backcrosses were released into two natural lakes. Parentage analysis using microsatellite loci facilitated the identification of survivors, so fitness was estimated in nature from the first-feeding stage. Results indicated that under certain conditions, domesticated fish survived at least as well as wild fish within the same environment. Relative growth and survival of the crosses, however, were highly dependent on environment. During the first summer, fastest-growing crosses had the highest survival, but this trend was reversed after one winter and another summer. Although the F1 hybrids showed evidence of outbreeding depression because of the disruption of local adaptation, there was little evidence of outbreeding depression in the backcrosses, and the second-generation backcrosses exhibited a wild-type phenotype. This information is relevant for assessing the multigenerational risk of escaped or released domesticated fish should they successfully interbreed with wild populations and provides information on how to minimize detrimental impacts of a conservation breeding and/or management programme. These data also further understanding of the selection pressures in nature that maintain submaximal rates of growth.

How the ladybird got its spots : effects of resource limitation on the honesty of aposematic signals

Prey species often possess defences (e.g. toxins) coupled with warning signals (i.e. aposematism). There is growing evidence that the expression of aposematic signals often varies within species and correlates with the strength of chemical defences. This has led to the speculation that such signals may be 'honest', with signal reliability ensured by the costliness of producing or maintaining aposematic traits. We reared larval seven-spot ladybirds (Coccinella septempunctata) on a Low or High aphid diet and measured the effects on warning signal expression (elytral carotenoid pigmentation, conspicuousness, spot size), levels of defensive alkaloids (precoccinelline, coccinelline), and relationships between these traits. High-diet individuals had greater total precoccinelline levels, and elytra carotenoid concentrations at adulthood which was detectable to a typical avian predator. However, larval diet did not significantly affect adult body mass or size, spot size or coccinelline levels. Elytra carotenoid concentrations correlated positively with total precoccinelline levels in both diet groups and sexes. However, the relationship between elytra carotenoid concentrations and total levels of coccinelline depended on sex: in both diet groups, elytra carotenoids and coccinelline levels were positively correlated in females, but negatively correlated in males. Spot size and coccinelline levels correlated positively in Low-diet individuals, but negatively in High-diet individuals. These results point to physiological linkages between components of aposematism, which are modulated by resource (i.e. food) availability and affect the honesty of signals. Developmental diet, but also sex, influenced the relationships between signals and toxin levels. Ladybirds are sexually size dimorphic, and thus in comparison with males, females may be more susceptible to resource limitation and more likely to be honest signallers. © 2012 The Authors. Functional Ecology © 2012 British Ecological Society.

Phenological response of sea turtles to environmental variation across a species' northern range

Variations in environmental parameters (e. g. temperature) that form part of global climate change have been associated with shifts in the timing of seasonal events for a broad range of organisms. Most studies evaluating such phenological shifts of individual taxa have focused on a limited number of locations, making it difficult to assess how such shifts vary regionally across a species range. Here, by using 1445 records of the date of first nesting for loggerhead sea turtles (Caretta caretta) at different breeding sites, on different continents and in different years across a broad latitudinal range (25-39 degrees ' N), we demonstrate that the gradient of the relationship between temperature and the date of first breeding is steeper at higher latitudes, i.e. the phenological responses to temperature appear strongest at the poleward range limit. These findings support the hypothesis that biological changes in response to climate change will be most acute at the poleward range limits and are in accordance with the predictions of MacArthur's hypothesis that poleward range limit for species range is environmentally limited. Our findings imply that the poleward populations of loggerheads are more sensitive to climate variations and thus they might display the impacts of climate change sooner and more prominently.

Climate-induced reaction norms for life-history traits in pythons

Climate change modelers predict increasingly frequent “extreme events,” so it is critical to quantify whether organismal responses (such as reproductive output) measured over the range of usual climatic conditions can predict responses under more extreme conditions. In a 20-year field study on water pythons (Liasis fuscus), we quantified the effects of climatically driven annual variation in food supply on demographic traits of female pythons (feeding rate, body size, body mass, and reproductive output). Reaction norms linking food supply to feeding rates and residual body mass were broadly linear, whereas norms linking food supply to female body size became curvilinear when a dramatic (flooding-induced) famine reduced the mean body size at sexual maturity. Thus, the reaction norms recorded over 16 years of “normal” (albeit highly variable) climatic conditions gave little insight into the population's response to a more extreme nutritional crisis.

Individual (co)variation in thermal reaction norms of standard and maximal metabolic rates in wild-caught slimy salamanders

Standard metabolic rate (SMR) and maximal metabolic rate (MMR) are fundamental measures in ecology and evolution because they set the scope within which animals can perform activities that directly affect fitness. In ectotherms, both SMR and MMR are repeatable over time when measured at a single ambient temperature (Ta). Many ectotherms encounter variable Ta from day to day and over their lifetime, yet it is currently unknown whether individual differences hold across an ecologically relevant range of Ta (i.e. thermal repeatability; RT). Moreover, it is possible that thermal sensitivity of SMR and MMR are important individual attributes, and correlated with one another, but virtually nothing is known about this at present. We measured SMR and MMR across an ecologically relevant Ta gradient (i.e. from 10 to 25 °C) in wild-caught salamanders (Plethodon albagula) and found that RT was significant in both traits. SMR and MMR were also positively correlated, resulting in a lower RT in absolute and factorial aerobic scopes (AAS and FAS). We found significant individual differences in thermal sensitivity for both SMR and MMR, but not for AAS and FAS. The intercept (at Ta = 0 °C) and the slope of the thermal reaction norms were negatively correlated; individuals with low MR at low Ta had a higher thermal sensitivity. Finally, individuals with a high thermal sensitivity for SMR also had high thermal sensitivity for MMR. Our results suggest that natural selection occurring over variable Ta may efficiently target the overall level of - and thermal sensitivity in - SMR and MMR. However, this may not be the case for metabolic scopes, as the positive correlation between SMR and MMR, in addition to their combined changes in response to Ta, yielded little individual variation in AAS and FAS. Our results support the idea that organisms with low metabolism at low Ta have a high metabolic thermal sensitivity as a compensatory mechanism to benefit in periods of warmer environmental conditions. Hence, our study reveals the importance of considering within-individual variation in metabolism, as it may represent additional sources of adaptive (co)variation.

Prenatal acoustic communication programs offspring for high posthatching temperatures in a songbird.

In many species, embryos can perceive and learn external sounds. Yet, the possibility that parents may use these embryonic capacities to alter their offspring's developmental trajectories has not been considered. Here, we demonstrate that zebra finch parents acoustically signal high ambient temperatures (above 26°C) to their embryos. We show that exposure of embryos to these acoustic cues alone adaptively alters subsequent nestling begging and growth in response to nest temperature and influences individuals' reproductive success and thermal preferences as adults. These findings have implications for our understanding of maternal effects, phenotypic plasticity, developmental programming, and the adaptation of endothermic species to a warming world.

Effect of deformation schedule on the microstructure and mechanical properties of a thermomechanically processed C-Mn-Si transformation-induced plasticity steel

Thermomechanical processing simulations were performed using a hot-torsion machine, in order to develop a comprehensive understanding of the effect of severe deformation in the recrystallized and nonrecrystallized austenite regions on the microstructural evolution and mechanical properties of the 0.2 wt pct C-1.55 wt pct Mn-1.5 wt pct Si transformation-induced plasticity (TRIP) steel. The deformation schedule affected all constituents (polygonal ferrite, bainite in different morphologies, retained austenite, and martensite) of the multiphased TRIP steel microstructure. The complex relationships between the volume fraction of the retained austenite, the morphology and distribution of all phases present in the microstructure, and the mechanical properties of TRIP steel were revealed. The bainite morphology had a more pronounced effect on the mechanical behavior than the refinement of the microstructure. The improvement of the mechanical properties of TRIP steel was achieved by variation of the volume fraction of the retained austenite rather than the overall refinement of the microstructure.

Effect of microstructure on the stability of retained austenite in transformation-induced-plasticity steels

Two Fe-0.2C-1.55Mn-1.5Si (in wt pet) steels, with and without the addition of 0.039Nb (in wt pet), were studied using laboratory rolling-mill simulations of controlled thermomechanical processing. The microstructures of all samples were characterized by optical metallography, X-ray diffraction (XRD), and transmission electron microscopy (TEM). The microstructural behavior of phases under applied strain was studied using a heat-tinting technique. Despite the similarity in the microstructures of the two steels (equal amounts of polygonal ferrite, carbide-free bainite, and retained austenite), the mechanical properties were different. The mechanical properties of these transformation-induced-plasticity (TRIP) steels depended not only on the individual behavior of all these phases, but also on the interaction between the phases during deformation. The polygonal ferrite and bainite of the C-Mn-Si steel contributed to the elongation more than these phases in the C-Mn-Si-Nb-steel. The stability of retained austenite depends on its location within the microstructure, the morphology of the bainite, and its interaction with other phases during straining. Granular bainite was the bainite morphology that provided the optimum stability of the retained austenite.