Differentiated phenotypic plasticity in larvae of the cannibalistic salamander Hynobius retardatus

Alternative phenotypes in natural populations can arise from either genetic polymorphism or an environmentally induced phenotype, that is, polyphenism. Evolutionary models of polyphenism developed by theoretical studies predict that polyphenism is favored when there are environment-dependent fitness trade-offs between alternatives and that the threshold frequency for a facultative switch between alternative phenotypes is adjusted in accordance with different selection regimes. The broad-headed (alternative) larval morph of Hynobius retardatus, which is induced by crowding with conspecifics or heterospecific anuran (Rana pirica) larvae, is a representative example of cannibalistic polyphenism. Morph induction by such proximate factors must reflect evolutionary (conditional frequency-dependent) processes. To clarify the role of frequency-dependent processes in polyphenism, I investigated the occurrence rate of the broad-headed morph under experimental crowding conditions (low conspecific, high conspecific, and high heterospecific densities) using larvae from eight natural populations with different larval densities of conspecifics and heterospecifics, and found interpopulational differences in the expression of the morph. Thus, there is a larval density-dependent equilibrium frequency of the morph in each pond, suggesting that the local switch point for morph induction was modified by selection to produce evolved differences between ponds. The evolution of such interpond differences has three necessary conditions: (1) There are pond-dependent fitness trade-offs between alternatives, (2) The maintenance of the morph is costly, and (3) The presence of conspecific or, especially, heterospecific larvae provides a reliable cue to the receiver.     

Morphological variation and phenotypic plasticity of buoyancy in the macroalga Turbinaria ornata across a barrier reef

Many aspects of morphology of benthic algae (length, surface area-to-volume ratio, and blade undulation) are plastic traits that vary in response to physical factors (such as light or water flow environment). This study examines whether frond buoyancy is a plastic trait, and whether differences in morphology including buoyancy affect the potential persistence of macroalgae in habitats characterized by different water flow regimes. Fronds of the tropical alga Turbinaria ornata in protected backreef environments in Moorea, French Polynesia possess pneumatocysts (gas-filled floats) and experience positive buoyant forces, whereas fronds in wave-exposed forereef sites either lack pneumatocysts entirely or have very small, rudimentary pneumatocysts and experience negative buoyant forces. Forereef fronds transplanted to the backreef developed pneumatocysts and experienced increased buoyant force indicating that buoyancy is a phenotypically plastic trait in T. ornata. In comparing the potential for dislodgement by drag, drag was greater on forereef fronds at low flow speeds as these fronds were stiffer and did not bend over at low flow speeds and therefore were less streamlined in the flow than backreef algae, which bent easily. The environmental stress factor (ESF) (a measure of the likelihood of detachment for a frond in its habitat) was higher for forereef than backreef fronds at all flow speeds. When examined with respect to the flow velocities likely in their respective habitats however, the chance of detachment for backreef and forereef was similar. Neither backreef nor forereef fronds were predicted to break under normal, non-storm conditions, but both were predicted to break in storms. Strong forereef morphologies are well suited to habitats characterized by rapid flow, whereas the weaker, buoyant, tall backreef fronds are well suited to habitats where crowding and shading is common but hydrodynamic forces are low.     

Population structure in the coral Pavona cactus: clonal genotypes show little phenotypic plasticity

Electrophoretic data were used to examine the relationship between genotype and growth form, and to assess the contribution of asexual reproduction to recruitment within six local populations of the agaricid coral Pavona cactus from the central and northern Great Barrier Reef. The data revealed the presence of highly replicated clonal genotypes in the five densest populations. In three cases, samples of 50 to 60 colonies collected from 25 m² areas within the Eclipse Island and Pandora Reef populations each consisted of colonies with only two distinct 4-locus genotypes. More intensive sampling of the population at Eclipse Island showed that colonies with the same 4-locus genotypes were separated by distances of up to 93 m. In contrast, the population at Watson's Bay (Lizard Island) consisted of a few widely scattered and genetically distinct colonies. The samples collected from each population contained a range of growth forms including, in some cases, the entire morphological range described for this species. A strong association of genotype and growth form was detected in samples from all populations, with the exception of Watson's Bay where no such comparison was possible. Nevertheless, some genotypes were represented by more than one growth form and this could reflect the effects of limited phenotypic plasticity. The effects of asexual reproduction reduced the value of these data as a test of the genetic connectedness of the six populations studied. The genetic distance between samples was not simply correlated with geographic distance. This may reflect either the true structure of the breeding population(s) or the effects of asexual reproduction on estimates of allelic frequencies.Contribution No. 47 from the Ecology and Genetics Group of the University of Wollongong     

Deterministic approach to the study of the interaction predator-prey in a chemostat with predator mutual interference. Implications for the paradox of enrichment

Our understanding of predator-prey systems has progressed in recent decades mainly due to the ability to test models in chemostats. This study aimed to develop a deterministic model using differential equations to reproduce the dynamics of the interaction of a predator and a prey in a two stage chemostat focusing in the proposed previous prey dependent model of Fussmann et al. (2000) [Fussmann, G.F., Ellner, S.P., Shertzer, K.W., Hairston Jr., N.G., 2000. Crossing the Hopf bifurcation in a live predator-prey system. Science 290, 1358-1360]. The main problem with that model, but parameterized with the values obtained in this study (particularly the concentration of nutrient), was that the temporal trajectory of both the prey and the predator showed very high peaks that eventually led to the extinction of predator in all cases. In the same way the experimental time series obtained in this study does not exhibit the behavior predicted by the model of Fussman et al. On the contrary, as prey density increases, the system actually becomes more stable. Finally, the model that best explained the behavior of the predator and prey in the chemostat, at medium to high dilution rates, was the ratio dependent (algae-nitrogen) model with mutual interference measured in the chemostat (rotifer-alga) and that incorporated the age structure of the predator. Qualitative analysis of the dynamic behavior enabled evaluation of coexistence at equilibrium, coexistence on limit cycles, extinction of the predator or extinction of both populations.     

Interactions between morphological and physiological plasticity optimize energy acquisition in corals

Morphological plasticity in response to environmental heterogeneity may be performance enhancing or may simply result from an intrinsic instability in morphology during development. Although patterns of morphological change are well documented for numerous taxa, it is often unclear whether this plasticity enhances the performance of organisms in the habitat to which they have acclimatized. Reef-building corals are an ideal model system in which to investigate this question. We here develop a three-dimensional geometric model and present a comprehensive photosynthesis data set with experimentally calibrated photosynthesis models that predicts energy acquisition by foliose corals as a function of colony shape. This allows us to assess the extent to which changes in colony morphology along an environmental gradient track the predicted optimal colony morphologies. Our results provide strong evidence that phenotypic plasticity in foliose corals optimizes photosynthetic energy acquisition and is not simply a mechanism to increase light capture. We show that the optimal morphology is constrained at the boundaries of the environmental gradient, with non-optimal morphologies in these habitats having greatly reduced energy acquisition. However, at the center of the environmental gradient, flexibility in photophysiology allows energy acquisition to be very similar for multiple morphologies. Our results highlight the importance of phenotypic plasticity at multiple scales. Variation in overall morphology is important at niche boundaries at which conditions are consistently more stressful, whereas physiological flexibility is important in intermediate and less predictable habitats in which a rapid and reversible response to environmental fluctuations is required.     

不同底质和透明度下马来眼子菜的表型可塑性研究

马来眼子菜(Potamogeton malaianusMiq.)是目前太湖沉水植物优势种之一。文章比较分析了3种不同底质和透明度情况下马来眼子菜的生长特征,探索其在太湖不断扩展的原因及其适应性。分别选取粘土质粉砂、粉砂和下蜀黄土底质上生长的马来眼子菜进行观察试验。结果表明,这3种底质在粒度组成、营养盐和分布特征上具有显著差异。在太湖粘土质粉砂和粉砂底质上马来眼子菜的生物量比下蜀黄土底质上的马来眼子菜高,其生物量随着底质营养盐的增加而增加。马来眼子菜与觅光相关的形态指标,如高度、节间距、节数、叶数、叶长及叶面积均随着底质营养盐的增加而显著增加。马来眼子菜的冠层高度与水体的透明度呈显著负相关,表明其对剧烈变化的湖泊光环境具有很强的适应能力。马来眼子菜在不同底质上的形态可塑性是其优先占据湖泊资源成为优势种的重要原因。     

The role of turbidity as a constraint on predator-prey interactions in aquatic environments

Many of the world's most productive aquatic ecosystems usually contain turbid water. Paradoxically, many fish species that live in these habitats are also those that often rely on vision to detect their predators and their prey. For these fish, turbidity will reduce the distance at which predator-prey interactions occur, and there should be a reduction in the opportunity for behavioural modification to control the risk of predation. Under these conditions, we predict that most antipredator behaviour will become ineffective and that predator-prey interactions in turbid water will be primarily characterised by the direct effects of predator consumption of prey, rather than behavioural modification reducing the growth rates of prey. This hypothesis was tested in two laboratory experiments. The first experiment investigated how water turbidity, risk of predation, and their interaction affect habitat choice decisions by fathead minnows (Pimephales promelas). These data demonstrate that fathead minnows reduce their use of dangerous habitats, but that this effect is diminished in turbid water. A second experiment examined mortality patterns when these fish were preyed upon by yellow perch (Perca flavescens) in clear and turbid water. Absolute mortality rates were unaffected by visibility, but patterns of mortality were random in turbid water and skewed towards smaller individuals in clear water. Combined, these results support our hypothesis and suggest that the impact of predation risk will be reduced in turbid aquatic ecosystems. Received: 23 May 1996 / Accepted after revision: 18 November 1996     

Autotomy shapes the trade-off between seeking cover and foraging in larval damselflies

Animals commonly choose between microhabitats that differ in foraging return and mortality hazard. I studied the influence of autotomy, the amputation of a body part, on the way larvae of the damselfly Lestes sponsa deal with the trade-off between foraging or seeking cover. Survival of Lestes larvae when confronted with the odonate predator Aeshna cyanea was higher in a complex than in a simple microhabitat, indicating that this more complex microhabitat was safer. Within the simple microhabitat, larvae without lamellae had a higher risk for mortality by predation than larvae with lamellae, showing a long-term cost of autotomy. When varying the foraging value (food present or absent) and predation risk (encaged predator or no predator) in the simple microhabitat, larvae with and without lamellae responded differentially to the imposed trade-off. All larvae spent more time in the simple microhabitat when food was present than when food was absent. Larvae without lamellae, however, only sporadically left the safe microhabitat, irrespective of the presence of the predator. In contrast, larvae with lamellae shifted more frequently towards the risky microhabitat than those without lamellae, and more often in the absence than in the presence of the predator. These decisions affected the foraging rates of the animals. I show for the first time that refuge use is higher after autotomy and that this is associated with the cost of reduced foraging success. The different microhabitat preferences for larvae with and without lamellae are consistent with their different vulnerabilities to predation and demonstrate the importance of intrinsic factors in establishing trade-offs. Received: 4 June 1999 / Received in revised form: 18 August 1999/ Accepted: 18 August 1999     

Hatching plasticity in a Southeast Asian tree frog

Hatching, the life history switch point between embryonic and larval or subadult stages, has traditionally been regarded as a fixed event in an organism’s development. This notion has been challenged by reports of environmentally cued hatching in recent years, which show embryos improve fitness by hatching in response to mortality risks. Here, we present evidence of accelerated hatching due to predation at two points during embryonic development in Chiromantis hansenae. Young embryos (0 day old) exposed to simulated predation hatched earlier compared to undisturbed clutches. Old embryos (4 days old) subjected to direct katydid predation had more immediate responses, hatching <1 h after predation on average. Hatching time was not correlated with female frog size, egg attendance time, or other predator cues. Results confirm predator-cued hatching in a new family of amphibians and support hatching plasticity being a widespread and potentially ancestral condition. We suggest mechanisms and ecological basis of cue transmission and response in C. hansenae and point out potential further research.     

Contest behaviour in the speckled wood butterfly (Pararge aegeria): seasonal phenotypic plasticity and the functional significance of flight performance

Although contemporary animal contest theory emphasises the importance of physical asymmetries in resolving disputes, such asymmetries do not obviously settle fights in all groups. Territorial male butterflies, for example, compete via elaborate non-contact aerial interactions in which success is determined by relative persistence. Prior research suggests that the resolution of these contests is not clearly related to physical variables such as body size or energy reserves. However, given that the contests involve elaborate aerial manoeuvres, one long-standing suggestion is that asymmetries in flight performance, and thus flight morphology, may be important. We addressed this hypothesis via a manipulative investigation into the biophysical correlates of contest success in the speckled wood butterfly, Pararge aegeria. This species possesses the ability for significant adaptive phenotypic plasticity in relevant flight morphological parameters. We took advantage of this plasticity to rear 90 individuals of markedly varying flight morphologies, which we then pitted against each other in a semi-controlled experimental fashion. Multiple logistic and lognormal analyses provided little evidence for the relevance of morphological parameters, including relative flight musculature, wing loading and wing aspect ratio (wing length relative to area), to the outcome and/or duration of experimental contests. Instead, we found a positive effect of age upon contest success. Given that ability for high acceleration is strongly linked to variation in these morphological parameters, our findings suggest that flight performance is not a strong determinant of resource-holding potential in this notably territorial butterfly.     

Ancient behaviors of larval amphibians in response to an emerging fungal pathogen, <Emphasis Type="Italic">Batrachochytrium dendrobatidis</Emphasis>

Behaviors have evolved in response to various selection pressures over evolutionary time. However, not all behaviors are adaptive. Some presumably “ancient” behaviors, persistent for millions of years, may be detrimental in the face of novel selection pressures in modern times. These pressures include a multitude of emerging infectious diseases which may be stimulated by environmental changes. We examined how a globally emerging amphibian pathogen, Batrachochytrium dendrobatidis (BD), affected two key evolutionarily persistent behaviors displayed by amphibian larvae: aggregation and thermoregulation. Larval aggregation behavior is often essential for foraging, thermoregulation, and antipredator defense, but varies among species. Thermoregulatory behavior speeds larval development in ephemeral habitats. Specifically, we examined whether aggregation and thermoregulatory behaviors changed when exposed to the BD pathogen in two species (Bufo boreas and Rana cascadae) whose larvae aggregate in nature. In laboratory choice tests, larvae of neither species avoided infected conspecifics. BD-exposed B. boreas larvae aggregated, while unexposed R. cascadae larvae associated more frequently with BD-exposed conspecifics. There was no evidence of behavioral fever or altered thermoregulation in larvae of four species we examined (Pseudacris regilla, Rana aurora, B. boreas, R. cascadae). The absence of behavioral fever may suggest an inability of the larvae of some host species to mediate infection risk by this pathogen. Thermoregulatory behaviors may exhibit a high degree of evolutionary inertia in amphibian hosts because they are linked with host physiology and developmental rates, while altered aggregation behaviors could potentially elevate pathogen transmission rates, leading to increased infection risk in social amphibian species.     

土壤中乐果与微生物的相互作用研究

采用室内模拟实验,研究了土壤中乐果与微生物的相互作用.结果表明:(1)50、500、5 000 mg·kg-1的乐果在灭菌土中的降解速度十分缓慢,加药后15d,50 mg·kg-1的降解率为69.3%,500 mg·kg-1的降解率为26.7%,5 000 mg·kg-1的降解率为19.6%.在施过乐果的土壤中的降解速度非常迅速,加药后15d,50 mg·kg-1的降解率为99.7%,500 mg·kg-1的降解率为78.4%,5 000 mg·kg-1的降解率为54.6%.在未施过乐果的土壤中的降解速度则介于二者之间.说明微生物在乐果的降解中起着非常重要的作用,同时,施药可以诱发乐果降解菌的生成从而加速乐果的降解.(2)在各处理的土壤中,50 mg·k-1的乐果对细菌数星有一定的刺激作用,500、5 000 mg·kg-1的乐果抑制细菌的生长.且随质量浓度的升高,抑制作用增强.但随加药时间的延长,又有所恢复.3种不同质量浓度的乐果对土壤真菌和放线菌种群数量的影响均表现为明显的抑制作用,且随药剂质量浓度的提高和加药时间的延长抑制趋势越明显.因此,可以选择真菌和放线菌作为土壤受乐果污染的敏感指示菌.(3)在同样处理条件下,施过乐果土壤中的微生物种群数量并不比未施过乐果土壤中的占优势,表明乐果在施过药的土壤中降解速度的加快并不是由微生物的数量决定的,而是由它们的降解能力决定的.     

Correlations between weather and recruitment of Nassarius pauperatus,an intertidal gastropod with a planktotrophic larval stage

Recruitment of the intertidal gastropod Nassarius pauperatus (Lamarck) was estimated annually for more than a decade at three sites in South Australia. Recruitment varied temporally, but was relatively synchronous amongst sites, even though these were at least 10 km apart. The density of recruits at each site was negatively correlated with evaporation and also with several indices of temperature for February. We postulate that either high temperatures or desiccation cause the death of recently settled individuals, which may be unable to avoid these extremes by burrowing deeply into the substratum.     

Experimental evidence of specialised phenotypic roles in a mobbing raptor

Group living is associated with costs but also with potential benefits, such as a decrease in predation risk through, for example, higher defence efficiency. Mobbing is among the most specialised forms of anti-predator strategies involving group defence and has mainly been investigated in passerine birds and some mammals. Variation in the mobbing response has been found in several species according to phenotypic variation such as sex or age. This suggests that there are differential benefits between mobbers, which may have promoted individual specialisation in mobbing behaviour. We studied mobbing behaviour in a communal roosting raptor, the Marsh harrier (Circus aeruginosus), which shows active group defence. Our study population exhibits extreme colour polymorphism, with two colour morphs in males, as well as sexual dichromatism and colour variation with age. We used different decoys, placed at different distances from the roost, to manipulate experimentally the perceived predation risk and to elicit mobbing behaviour. Using the experimental design that maximised mobbing response in harriers, we then focused on the sequence and the specific behaviours involved in recruitment of mobbers, and whether individual investment in terms of defence was associated with phenotypic characteristics of individuals (i.e. sex, age and colour morph). We found that the main behaviour involved in successfully attracting mobbers was alarm calling. We also detected differential individual investment in relation to sex and age, but more importantly, we provide the first evidence for specialised male phenotypic roles during mobbing events, signalled by colour polymorphism: grey males tended not to be involved in mobbing and almost never behaved as recruiters or mobbers, while brown males behaved mainly as recruiter birds. These findings suggest that colour morph may signal the individual’s anti-predatory abilities through different behavioural strategies between males.     

Temperature cues gametogenesis and larval release in a tropical sponge

Determining the reproductive processes of benthic invertebrates is central to our understanding of their recruitment and population dynamics. Sexual reproduction of the gonochoric and viviparous Great Barrier Reef sponge, Luffariella variabilis (Poléjaeff 1884) was quantified from histological samples collected over two reproductive seasons (2004 and 2005). Gametogenesis commenced for females at a water temperature of 21°C, the lowest water temperature of the year. Spermatogenesis occurred above 22.5°C with sperm asynchronously developed and released from August or September to October. Oocytes developed asynchronously from July to September, embryos from September to December, and larvae from November to December. Female reproduction terminated in December (after larval release) prior to the highest mean annual water temperature of 30°C in January. There was a significant (35%) decrease in female reproductive output in 2005 compared to 2004, as measured by the reproductive index (0.68 ± 0.12 female reproductive propagules mm⁻² of mesohyl in 2005 compared with 1.05 ± 0.10 mm⁻² in 2004). This corresponded with delayed oogenesis and spermatogenesis, and a shortened larval development cycle corresponding with a delayed minimum temperature (21°C) in August of 2005 compared with July 2004. Accordingly, the maximum percentage of the mesohyl occupied by female reproductive propagules (eggs, embryos and larvae) was also reduced by 60% in 2005 (overall mean of 13.04% in October 2004 compared with 5.35% in October 2005). However, the mean sizes of individual female propagules remained the same from year to year. Males in contrast, showed no overall difference in either reproductive index or percentage occupation of the mesohyl between 2004 and 2005. The lowered reproductive output (∼35%) of females of L. variabilis associated with delayed minimum water temperatures may have important implications for population reproductive success where oogenesis and spermatogenesis and larval release are cued by minimum and maximum water temperatures, respectively.     

Orientation ofRhizophagus grandis (Coleoptera: Rhizophagidae) to oxygenated monoterpenes in a species-specific predator-prey relationship

Summary The predator,Rhizophagus grandis, is linked to its specific prey,Dendroctonus micans, by semiochemical signals regulating oviposition and long range orientation. A mixture of simple oxygenated monoterpenes [(–)—fenchone, (–)—pinocamphone, rac. camphor, terpinene-4-ol, borneol, fenchol and verbenone], identified from the frass ofD. micans has been found to be extremely active in a flight windtunnel as a long—range attractant for the predator. The mixture elicited 84 % of the response to larval frass of the prey. Excluding pinocamphone from the mixture did not influence its activity. Also, changing the absolute configurations of some of the components (fenchol, terpinene-4-ol and borneol) did not influence the predators' response. However, the addition of (–)—-terpineol increased the attractivity of the synthetic blend to almost that of larval frass of the prey. Ecological implications of the identified semiochemicals and their use in pest management are discussed.