The defensive behavior of the edible snail]

Literary and experimental data are summarized on organization of defensive behaviour in Helix. Muscular and neuronal mechanisms participating in this form of behaviour are reviewed.     

Crayfish induce a defensive shell shape in a freshwater snail

Abstract. This study was undertaken to determine whether intraspecific variation in shell morphology of the freshwater snail Elimia (= Goniobasis ) livescens is caused by predator-induced morphological changes. Juvenile snails from 3 populations were grown in the presence of effluent from predatory crayfish feeding on conspecific snails or in effluent from conspecific snails only. Snails from one population, Clear Creek, exhibited a predator-induced morphology; they grew a narrow body whorl when exposed to the effluent from crayfish. Experimental feeding trials with crayfish and snails from Clear Creek were conducted to determine whether a narrow body whorl reduced predation in the presence of the crayfish Orconectes propinquus . In the feeding trials, snails with a narrow body whorl were eaten less frequently than snails with wider shells. However, there was no difference in overall size (length) between snails that were eaten and those that were not. Thus, juvenile snails from Clear Creek exposed to the presence of crayfish were induced to develop relatively narrow shells, which reduce the risk of successful attack by these crayfish. Hence the induced shell morphology is probably a defense against predation by crayfish.     

Age-related changes in defensive traits of Acacia tortilis Hayne

The theory of plant defences proposes that investments in physical and chemical defences are driven by the risk of herbivore damage, and limited by the cost of producing the particular defensive trait in terms of resources that could be directed to other sinks, such as growth and reproduction. We sampled twigs of 18 mature Acacia tortilis trees and their cohort of juveniles to test some predictions of this hypothesis. We expected a higher allocation of defensive traits to leaves and twigs in the young plants than in the mature ones as a result of a higher risk of damage by ungulates at the juvenile stage. Our results show that the juvenile plants produce more spines along their twigs, but have lower concentrations of phenolic compounds in their leaves than in the mature ones. We also expected a negative relation between the concentration of foliar nutrients and phenolic compounds, as predicted by the carbon/nutrient hypothesis. Only mature plants showed this pattern. Reproduction (in mature plants) and water stress (in juvenile plants) did not relate to allocation to secondary compounds as predicted by current hypotheses of plant defence.     

Electrophysiological study of 5,6-dihydroxytryptamine effect on defensive reflex conditioning in the snail

The role of serotonin in expression of membrane properties of identified neurons was studied during defensive reflex conditioning using the neurotoxic analogue of serotonin 5,6-dihydroxytryptamine (5,6-DHT). The defensive reflex conditioning in snails was destroyed on the second day after second injection of 5,6-DHT. Through the 1st weeks after second injection of 5,6-DHT the snails were learned but worse than snails after injection of saline solution. This result shows the recovery of snail's learning ability within 2 weeks after the second injection of 5,6-DHT. It was found that injection of 5,6-DHT prevented the decrease of membrane and threshold potentials of command neurons during defensive reflex conditioning as compared with the snails injected with 5,6-DHT without learning.     

Long-term evolution of multilocus traits

 We analyze monomorphic equilibria of long-term evolution for one or two continuous traits, controlled by an arbitrary number of autosomal loci and subject to constant viability selection. It turns out that fitness maximization always obtains at long term equilibria, but in the case of two traits, linkage determines the precise nature of the fitness measure that is maximized. We then consider local convergence to long term equilibria, for two multilocus traits subject to either constant or frequency dependent selection. From a model of long-term dynamics near an equilibrium we derive a criterion of local long-term stability for 2-dimensional equilibria. It turns out that mutation can be a decisive factor for stability. Received 26 January 1994; received in revised form 26 September 1994     

Using knockout mutants to reveal the growth costs of defensive traits

We used a selection of Arabidopsis thaliana mutants with knockouts in defence genes to demonstrate growth costs of trichome development and glucosinolate production. Four of the seven defence mutants had significantly higher size-standardized growth rates (SGRs) than the wild-type in early life, although this benefit declined as plants grew larger. SGR is known to be a good predictor of success under high-density conditions, and we confirmed that mutants with higher growth rates had a large advantage when grown in competition. Despite the lack of differences in flowering-time genes, the mutants differed in flowering time, a trait that strongly correlated with early growth rate. Aphid herbivory decreased plant growth rate and increased flowering time, and aphid population growth rate was closely coupled to the growth rate of the host plant. Small differences in early SGR thus had cascading effects on both flowering time and herbivore populations.     

Effects of temperature on respiration defensive behavior and locomotion of fresh water snail Lymnaea stagnalis

Temperature dependence of lung respiration, defensive behavior and locomotion of Lymnaea stagnalis snail was studied. At the temperature in the range of 4-6 degrees C the rates of locomotion and respiration were reduced (as compared to control temperature of 14-16 degrees C), whereas defensive reactions were much more intense. Vice versa, the temperature rise to 24-26 degrees C activates respiration and locomotion but inhibits defensive behavior. It is suggested that the observed changes in Lymnaea behavior result from temperature-dependent reactions of neurons underlying these activities.     

The role of dopamine and serotonin in modulating the defensive behavior of the edible snail

Dopamine application in concentration of 10(-5)-10(-6) M into saline around the snail CNS leads to decrease of excitability of LPa7 neurone which is presynaptic in relation to defensive behaviour command neurones, and to decrease of amplitude of monosynaptic excitatory postsynaptic potential (EPSP) in the command neurones elicited by intracellular stimulation of LPa7 neurone. Besides, the dopamine causes a decrease of summated EPSP amplitude in the studied neurones in response to intestinal nerve stimulation (70% in average), a change of rest potential towards hyperpolarization for 6-8 mV, a reduction of the command neurones input resistance (20% in average). The described influences can lead to a general increase of the threshold of defensive system reaction to stimulation. Dopamine action on the defensive behaviour command neurones is significantly weakened in serotonine presence. Against the dopamine background, the efficiency of serotonine influence on the value of EPSP in command neurones in response to testing stimulus is reduced. According to the obtained data, a conclusion is made that interrelation of dopamine and serotonine concentrations can be a base for formation of behaviour choice in snail.     

Combined use of two defensive traits in pupae of Scymnus posticalis ladybirds

Coccinellid pupae have various defensive traits, such as keeping their final‐instar larval exuviae and secreting droplets containing defensive chemicals at the tips of glandular hairs. Although each of these traits has been described separately in different species, it is unclear how each defensive trait functions when more than one trait coexists. We found that pupae of the ladybird Scymnus posticalis have two different types of traits; they were partially covered with their own final‐instar larval exuviae, with thick wax structures, and had many glandular hairs secreting transparent droplets on the exposed parts of the body without waxy exuviae (head and medial parts of thoracic and abdominal dorsum). In this study, the defensive effects of these two traits were evaluated against two ant species, Tetramorium tsushimae and Lasius japonicus, which are potential predators. When encountering a pupa, neither ant species were able to make frequent physical contact with the exposed parts of the pupa because of the presence of the waxy exuviae. When touching the exposed body part that was covered with droplets, both ant species immediately stepped backward and then swept their antennae. The droplets suppressed ant feeding when mixed with sucrose solution, suggesting that the droplets contain repellent chemicals. These results indicate that the two traits of S. posticalis pupae (the larval waxy exuviae and the droplets) effectively play defensive roles against ants, functioning as a physical obstruction and a chemical repellent, respectively.     

抗烟草黄瓜花叶病毒多糖筛选及其对烟草防御酶活性的影响

[目的]筛选出对烟草黄瓜花叶病毒病有良好抑制作用的多糖并探索其对烟叶防御酶活性的影响.[方法]采用半叶法,测定了安络小皮伞多糖等21种真菌多糖在枯斑三生烟上对CMV的钝化、预防及治疗效果,并测定了抗病毒多糖处理后普通烟NC-89体内防御酶的变化.[结果]安络小皮伞多糖对CMV具有较好的钝化及预防效果,其200倍液与等量供试病毒液混合30 min后接种,钝化效果为83.41%;喷施安络小皮伞多糖200倍液24 h后接毒处理,预防效果可高达93.15%.安络小皮伞多糖对CMV防治机理的研究表明,多糖处理后烟草相关防御酶POD、PAL和PPO活性增强,其中喷施安络小皮伞多糖24 h后接毒处理的酶活增加最为显著,该处理烟苗的POD、PAL和PPO的酶活峰值分别可增加至对照的2.74、3.45和2.82倍.[结论]安络小皮伞多糖通过增强烟草体内防御酶活性而提高烟草对烟草黄瓜花叶病毒病的抗性.     

Underdispersion and overdispersion of traits in terrestrial snail communities on islands

Understanding and disentangling different processes underlying the assembly and diversity of communities remains a key challenge in ecology. Species can assemble into communities either randomly or due to deterministic processes. Deterministic assembly leads to species being more similar (underdispersed) or more different (overdispersed) in certain traits than would be expected by chance. However, the relative importance of those processes is not well understood for many organisms, including terrestrial invertebrates. Based on knowledge of a broad range of species traits, we tested for the presence of trait underdispersion (indicating dispersal or environmental filtering) and trait overdispersion (indicating niche partitioning) and their relative importance in explaining land snail community composition on lake islands. The analysis of community assembly was performed using a functional diversity index (Rao's quadratic entropy) in combination with a null model approach. Regression analysis with the effect sizes of the assembly tests and environmental variables gave information on the strength of under‐ and overdispersion along environmental gradients. Additionally, we examined the link between community weighted mean trait values and environmental variables using a CWM‐RDA. We found both trait underdispersion and trait overdispersion, but underdispersion (eight traits) was more frequently detected than overdispersion (two traits). Underdispersion was related to four environmental variables (tree cover, habitat diversity, productivity of ground vegetation, and location on an esker ridge). Our results show clear evidence for underdispersion in traits driven by environmental filtering, but no clear evidence for dispersal filtering. We did not find evidence for overdispersion of traits due to diet or body size, but overdispersion in shell shape may indicate niche differentiation between snail species driven by small‐scale habitat heterogeneity. The use of species traits enabled us to identify key traits involved in snail community assembly and to detect the simultaneous occurrence of trait underdispersion and overdispersion.     

Microgeographic evolution of snail shell shape and predator behavior

Genetic divergence in geographically isolated populations is a prerequisite for allopatric speciation, one of the most common modes of speciation. In ecologically equivalent populations existing within a small, environmentally homogeneous area, an important role for environmentally neutral divergence is often found or inferred. We studied a species complex of conspicuously shaped Opisthostoma land snails on scattered limestone outcrops within a small area of lowland rainforest in Borneo. We used shell morphometrics, mitochondrial and nuclear DNA sequences, and marks of predation to study the factors involved in allopatric divergence. We found that a striking geographic divergence exists in shell morphology, which is partly associated with neutral genetic divergence. We also found geographic differentiation in the behavior of the snails' invertebrate predator and evidence of an evolutionary interaction between aspects of shell shape and predator behavior. Our study shows that adaptation to biotic aspects of the environment may play a more important role in allopatric speciation than previously suspected, even on a geographically very small scale.     

The evolution of group-level pathogenic traits

A group-selection model for the evolutionary origin of phase-variation in E. coli is proposed. Populations of commensal strains of E. coli populating mammalian hosts modulate its immune defenses through population-level control of the expression of fimbriae. At any time only a proportion of the population expresses these cell-surface adhesins. Collectively they elicit a host-based nutrient release if the fimbriae expression is low. Too high levels of fimbriation would provoke an inflammatory response and thus intolerable conditions for the cells. The optimal level of fimbriation is a group property and its evolution is difficult to explain by naive individual selection scenarios. This article presents a computational model to simulate the evolution of fimbriae. The two main conclusions of this contribution are: (i) the evolution of this group property requires the population to be partitioned into weakly interacting sub-populations. (ii) Given certain scenarios evolution consistently under-performs, in the sense that it does not find the optimal level of fimbriation.     

Electrophysiological study of effects of chronic injection of caffeine on defensive reflex conditioning in grape snail

It was found that chronic injection of caffeine to grape snail increases a velocity of elaboration of conditioned defensive reflex. It was shown that after daily injection of caffeine immediately after procedure of learning the conditioned defensive reflex elaborated faster than daily injection before procedure of learning. It has been shown, that chronic injection of caffeine both in naive as well as learned snails led to depolarizing shift of membrane potential and to decrease of threshold potential of command neurons of the defensive behaviour of grape snails. It was also found that addition of caffeine in bath solution led to decrease of threshold of generation of action potential of command neurons both in intact and learned snails. The resting membrane potential of command neurons was not changed.     

Behavioral changes during the acquisition of long-term sensitization of the defensive reflex in the edible snail

Reaction of long-term sensitization (LTS) of defensive reflex was elaborated in the snail. It was accompanied by a considerable increase of the time of closing of pneumostome. The results show that LTS is preserved for more than two week and may be a model of long-term memory.     

Investment in boney defensive traits alters organismal stoichiometry and excretion in fish

Oecologia - Understanding how trait diversification alters ecosystem processes is an important goal for ecological and evolutionary studies. Ecological stoichiometry provides a framework for...     

Co‐occurrence of defensive traits in the potato aphid Macrosiphum euphorbiae

1. Insecticide usage selects strongly for resistance in aphid populations, but this could entail fitness costs in other resistance traits. The potato aphid Macrosiphum euphorbiae Thomas exhibits intraspecific variation in susceptibility to parasitism by braconid wasps and provides a suitable species to study the relation between the defensive traits of parasitism and insecticide resistance. 2. Clonal lines (23 in total) of M. euphorbiae were established from aphids collected in 2013 from geographically separate populations in the U.K. Clonal lines belonged to five aphid genotypes, but one genotype predominated (78% of samples), and the facultative endosymbiont Hamiltonella defensa was detected in c. 40% of lines. 3. Total esterase activity in aphid tissues varied significantly between aphid genotypes and collection areas, but there was no clear pattern in relation to H. defensa infection or between collection sites likely to differ in insecticide pressures. 4. Five clonal lines representing low to moderate levels of enzyme activity, which included different aphid genotypes and presence/absence of H. defensa infection, were assayed for their susceptibility to the parasitoid wasp Aphidius ervi Haliday. Aphid mummification varied significantly between aphid genotypes, with low values in one genotype of aphids irrespective of H. defensa presence. 5. The results revealed that aphid lines belonging to the parasitism‐resistant genotype exhibited moderate levels of total esterase activity, indicating a competitve advantage for this genotype of M. euphorbiae when exposed to chemical and biological control factors in agroecosystems.     

Nonrandom variation of morphological traits across environmental gradients in a land snail

Morphological variation is often attributed to differential adaptations to diverse habitats, but adaptations to a similar environment do not necessarily result in similar phenotypes. Adaptations for water and heat budget are crucial for organisms living in arid habitats, and in snails, variation in shell morphology has been frequently attributed to selection by stressful environmental factors. However, their phenotypic divergence often is not accompanied by a relevant niche differentiation and consistent relationships with environmental correlates are lacking. In the pulmonate genus Albinaria, there is great size and shape variation between and within species, and there are two major shell sculpture morphotypes, ribbed and smooth. We used 62 populations of 28 Albinaria species, taking into account their phylogeny, to examine the variation of shell traits (sculpture, size, shape), their effect on water and heat budget, and their association with geographical and climatic gradients. We found unambiguous size and shape discrimination between the two morphotypes. Ribbed shells are lighter, taller, and slimmer and have a smaller aperture than the smooth ones. Moreover, significant correlations between shell traits and heat/moisture budget and climate/geography were revealed. Ribbed and taller shells retain more water on their shell surface, and on the other hand, smooth shells exhibit lower water permeability. Therefore, two strategies are being used to prevent water loss, active retention or resistance to loss. Consequently, different alternative solutions evolved and were retained as responses to the same stressful factor by the two distinct shell morphotypes. Larger shells occur in southern latitudes, mostly on islands, and at sites where there is a shortage of rainfall. Therefore, the variation of the examined traits is nonrandom with respect to location and to climate and their evolution can be attributed to selection by environmental factors, with water availability being the key driving agent of body-size variation.     

Rapid evolution of novel traits in microorganisms

The use of natural microorganisms in biotransformations is frequently constrained by their limited tolerance to the high concentrations of metabolites and solvents required for effective industrial production. In many cases, more robust strains have to be generated by random mutagenesis and selection. This process of directed evolution can be accelerated in mutator strains, which carry defects in one or more of their DNA repair genes. However, in order to use mutator strains, it is essential to restore the normal low mutation rate of the selected organisms immediately after selection to prevent the accumulation of undesirable spontaneous mutations. To enable this process, we constructed temperature-sensitive plasmids that temporarily increase the mutation frequency of their hosts by 20- to 4,000-fold. Under appropriate selection pressure, microorganisms transformed with mutator plasmids can be quickly evolved to exhibit new, complex traits. By using this approach, we were able to increase the tolerance of three bacterial strains to dimethylformamide by 10 to 20 g/liter during only two subsequent transfers. Subsequently, the evolved strains were returned to their normal low mutation rate by curing the cells of the mutator plasmids. Our results demonstrate a new and efficient method for rapid strain improvement based on in vivo mutagenesis.