Chronic alterations in serotonin function: dynamic neurochemical properties in agonistic behavior of the crayfish, Orconectes rusticus

The biogenic amine serotonin [5-hydroxytryptamine (5-HT)] has received considerable attention for its role in behavioral phenomena throughout a broad range of invertebrate and vertebrate taxa. Acute 5-HT infusion decreases the likelihood of crayfish to retreat from dominant opponents. The present study reports the biochemical and behavioral effects resulting from chronic treatment with 5-HT-modifying compounds delivered for up to 5 weeks via silastic tube implants. High performance liquid chromatography with electrochemical detection (HPLC-ED) confirmed that 5,7-dihydroxytryptamine (5,7-DHT) effectively reduced 5-HT in all central nervous system (CNS) areas, except brain, while a concurrent accumulation of the compound was observed in all tissues analyzed. Unexpectedly, two different rates of chronic 5-HT treatment did not increase levels of the amine in the CNS. Behaviorally, 5,7-DHT treated crayfish exhibited no significant differences in measures of aggression. Although treatment with 5-HT did not elevate 5-HT content in the CNS, infusion at a slow rate caused animals to escalate more quickly while 5-HT treatment at a faster rate resulted in slower escalation. 5,7-DHT is commonly used in behavioral pharmacology and the present findings suggest its biochemical properties should be more thoroughly examined. Moreover, the apparent presence of powerful compensatory mechanisms indicates our need to adopt an increasingly dynamic view of the serotonergic bases of behavior like crayfish aggression.     

Behavioral effects of serotonin and serotonin agonists in two crayfish species, Procambarus clarkii and Orconectes rusticus

Exogenous serotonin elicits several behaviors in Procambarus clarkii, including a flexed, elevated posture, reduced locomotion, and changes in aggressive behavior. We conducted experiments to determine if several serotonin agonists mimicked the behavioral effects of serotonin in two crayfish species, P. clarkii and Orconectes rusticus. Drugs tested were 1-(3-Chlorophenyl)-piperazine dihydrochloride (mCPP), Oxymetazoline, 2,5-dimethoxy-4-iodoamphetamine (DOI), CGS-12066A, and (+/-)-8-hydroxy-2-(di-n-dipropylamino) tetralin (8-OH-DPAT). In P. clarkii, mCPP most closely mimicked the effects of serotonin, significantly increasing the performance of the flexed, elevated posture and reducing locomotion; 8-OH-DPAT significantly reduced locomotion as well. Both of these drugs produced significant increases in elevated posture and decreases in locomotion in O. rusticus, and in this species, the drugs at test concentrations were more effective in eliciting these effects than serotonin. The effects of the drugs on behaviors performed during fighting bouts were variable. In both species, only 8-OH-DPAT significantly reduced several agonistic behaviors, and no agonist or 5-HT itself produced significant increases in agonistic behavior.     

Cytochemical observations of hemolymph cells during coagulation in the crayfish, Orconectes virilis

Smears of Orconectes virilis hemolymph were prepared by fixation in 10% formalin immediately upon withdrawal, and after an interval of 1, 5, or 15 minutes. These smears were then stained by a variety of histochemical means designed to identify a material which is released from both hyaline and granular cells during the coagulation process. Based upon reactions to various carbohydrate, lipid and protein tests, this material appears to consist of a glycoor muco-protein, and its localization and activity coincide with the ?fibrin ferment”? described by Hardy (1892). Its release is probably initiated from the hyaline cells, followed shortly thereafter by a similar activity of the granulocytes. While it cannot be ascertained by the methods used in this study whether or not this material participates actively in clotting, its release is nevertheless coincidental with coagulation.     

Histological and histochemical observations of the hemolymph cells in the crayfish, Orconectes virilis

Hemolymph cells of Orconectes virilis were stained during the months of August to November by a variety of histological and histochemical techniques. Cells were classified as hyaline cells, small granulocytes, and large granulocytes. Presence of mitochondrial enzymes was indicated by tests for succinic dehydrogenase and cytochrome oxidase. Reaction to test for the hydrolytic enzyme, leucine acylnaphthylamidase was intense in the granules of the large granulocytes. The PAS reaction indicated a mucopolysaccharide at the cell membrane. Lipid was found in all three hemolymph cell types of Orconectes virilis at the time of this study.     

Effects of fluoxetine on growth and behavior in the crayfish Orconectes rusticus

This laboratory study examined the effects of the specific serotonin reuptake inhibitor fluoxetine on growth following molting and on a range of behaviors in the crayfish Orconectes rusticus. For growth experiments, male Form I and Form II crayfish were weighed and measured and placed individually in water containing 0–500 μg/L of fluoxetine. They were held in fluoxetine or control water until they molted and were reweighed two weeks post-molt. In behavior experiments, juvenile and adult animals were held individually in 0, 2, 200, or 500 μg/L of fluoxetine for 10 days and tested in an open field arena to assess locomotion, thigmotaxis, sheltering, and habituation to a novel environment. Under our laboratory conditions, crayfish exposed to fluoxetine at 500 μg/L showed significantly enhanced growth: post-molt Form I animals had greater body weight and post-molt Form II animals had greater carapace length, relative to controls. In open field tests, juvenile crayfish exposed to 2 and 500 μg/L fluoxetine displayed significantly reduced locomotion compared to controls. The results indicate that crayfish growth and locomotion can be manipulated by short-term exposure to ambient fluoxetine, suggesting that this means of exposure may offer a useful and noninvasive way to examine drug effects in freely moving animals. However, effects were only observed at concentrations well above fluoxetine levels currently reported in the environment. This suggests that O. rusticus may be relatively resistant to this form of pharmaceutical pollution but whether effects would occur following long-term exposure to lower concentrations is unknown.     

Some factors regulating cohort production of the crayfish, Orconectes virilis

SUMMARY. Cohort production of the crayfish. Orconectes virilis , ranged from 148.8 to 70.2 kg in North Twin Lake (surface area 1.9 ha) and from 151.9 to 84.9 kg in West Lost Lake (surface area 1.4 ha). Both cohort production (P) and cohort mean biomass (B) showed a long-term decrease with time. The cohort P/B ratios did not show such a trend but remained steady. The P/B ratio approximated the life cycle instantaneous rate of growth (G) which could then be used for estimating cohort production from mean biomass.
Production remained consistently higher in West Lost Lake. Analysis of horizontal life table data revealed no inherent difference between survival and fecundity rates of the crayfish population in each lake. Slight differences in basin morphometry between the two lakes determined nursery habitat availability and this affected recruitment of young from each female brood stock. Small differences in carrying capacity between two similar sized nearby lakes, along with a decrease in carrying capacity over time, were sufficient to nullify attempts to manage these populations using a maximum sustainable yield model.     

The mate pursuit-predation avoidance tradeoff in the virile crayfish, Orconectes virilis

To better understand the tradeoff between mate pursuit and predation risk avoidance, the responses of male virile crayfish (Orconectes virilis) to female sex pheromone and conspecific alarm cue, alone and in combination were tested. Male crayfish were more active and exhibited less antipredator behavior in the pheromone treatment than in either the pheromone + alarm or alarm treatment. The results suggest a tradeoff in favor of risk avoidance when pheromone and risk stimuli are encountered together.     

The mate pursuit-predation avoidance tradeoff in the virile crayfish,Orconectes virilis

To better understand the tradeoff between mate pursuit and predation risk avoidance, the responses of male virile crayfish (Orconectes virilis) to female sex pheromone and conspecific alarm cue, alone and in combination were tested. Male crayfish were more active and exhibited less antipredator behavior in the pheromone treatment than in either the pheromone?+?alarm or alarm treatment. The results suggest a tradeoff in favor of risk avoidance when pheromone and risk stimuli are encountered together.     

Intercellular junctions of antennal gland epithelial cells in the crayfish,Orconectes virilis

Summary Labyrinth and nephridial canal cells of the crayfish (Orconectes virilis) antennal gland possess two types of intercellular junctions revealed by freeze-fracture studies. Apical margins of the cells are connected by long septate junctions. In replicas, these junctions consist of many parallel rows of 80–140 Å intramembrane particles situated on the PF membrane face (EF and PF fracture faces of Branton et al., 1975). Rows of pits are found on the EF fracture face and are deemed complementary to the rows of particles. Moreover, lateral margins of basal regions of the epithelial cells are attached by many intercellular junctions. These contacts are characterized in thin plastic sections by a narrow dense cytoplasmic plaque located subjacent to the plasma membrane at sites of adjoined cells, and 5 to 12 fine strands of dense material that extend across the intercellular gap between adjoined cells. In freeze-fracture replicas, EF intramembrane faces basal to the region of the plasma membrane containing septate junctions exhibit numerous discoid clusters of particles. The particle aggregates, assumed to represent freeze-cleave images of adhering junctions, range from 900 to 3,700 Å in diameter, with individual particles about 185 Å in diameter. These junctions appear to connect epithelial cell processes formed by basal infoldings of the plasma-lemma, and occur between adjacent cells as well as adjacent processes of a single cell. The discrete aggregates of particles resemble replicated desmosomes (Shienvold and Kelly, 1974) and hemi-desmosomes (Shivers, 1976); therefore, they probably do not constitute a basis for electrical coupling between antennal gland epithelial cells.Supported by the National Research Council of Canada     

A trematode parasite alters growth,feeding behavior,and demographic success of invasive rusty crayfish (Orconectes rusticus)

Nonindigenous species can cause major changes to community interactions and ecosystem processes. The strong impacts of these species are often attributed to their high demographic success. While the importance of enemy release in facilitating invasions has often been emphasized, few studies have addressed the role of parasites in the invasive range in controlling demographic success of potential invaders. Here we examine whether a trematode parasite (Microphallus spp.) can contribute to previously documented alternate states in the abundance of invasive rusty crayfish (Orconectes rusticus) in north temperate lakes in Wisconsin, USA. Microphallus infect O. rusticus after emerging from their first intermediate host, a hydrobiid snail. As previously documented, O. rusticus reduce densities of hydrobiid snails through direct predation and destruction of macrophyte habitat. Therefore, if Microphallus substantially reduce O. rusticus fitness, these parasites may reinforce a state of low crayfish abundance, and, at the other extreme, abundant crayfish may repress these parasites, reinforcing a state of high crayfish abundance. From samples collected from 109 sites in 16 lakes, we discovered (1) a positive relationship between crayfish infection intensity and hydrobiid snail abundance, (2) a negative relationship between parasite prevalence and crayfish abundance, and (3) a negative relationship between parasite prevalence and crayfish population growth. With experiments, we found that infection with Microphallus reduced foraging behavior and growth in O. rusticus, which may be the mechanisms responsible for the population reductions we observed. Overall results are consistent with the hypothesis that Microphallus contributes to alternate states in the abundance and impacts of O. rusticus.     

Exploration of potential microbial control agents for the invasive crayfish,Orconectes virilis

The introduced crayfish, Orconectes virilis (Cambarinae; Hagen 1870), has become a serious invasive species in Arizona (USA), altering stream ecosystems and contributing to the decline of native species. But because it is native to the eastern US, and related crayfish including endangered species inhabit nearby states, introduction of a biological control agent presents a unique challenge. This represents a feasibility study, in which we explored bacteria, nematodes, and a virus as potential biological control agents for O. virilis while avoiding harm to native species. White Spot Syndrome Virus (WSSV) from shrimp was found to be highly pathogenic and readily passed by cannibalistic behavior but not by water transmission. Potential activity of WSSV against representative non-target arthropods that may be found in the same ecosystems was also explored.     

The tradeoff between reproductive and food resources in the crayfish Orconectes virilis

The conflict between mating and foraging is one of the many tradeoffs associated with reproduction. The relative attraction to sex pheromone and food stimuli by male virile crayfish (Orconectes virilis) were tested using two experiments. In the first experiment, the responses by males to pheromone and food stimuli alone were assessed. In the second experiment, males were exposed to both pheromone and food stimuli in a choice maze. To determine the effects of feeding motivation, the males in the choice experiment were fed either one day or seven days prior to observation. Male crayfish exhibited similar responses to the pheromone and food stimuli individually. Males exhibited a preference for the pheromone stimulus over the food stimulus in the choice trials, and that preference that was not influenced by starvation.     

Bilateral and unilateral antennal lesions alter orientation abilities of the crayfish, Orconectes rusticus.

Numerous animals use chemical cues within their environments to execute various behaviors. One of these behaviors is orientation to an odor source. Crayfish, in particular, can orient to food sources under a number of different conditions. It has not been determined, however, what kind of search strategy these animals employ to successfully locate a food source. To determine the role of antennae and antennules in this behavior and to investigate different modes of orientation behavior, the orientation patterns of crayfish with complete and partial antennal lesions were examined. Detailed analysis of orientation paths confirmed that crayfish could not locate odor sources with either bilateral or unilateral lesions. This suggests that crayfish are using the spatial information obtained from these appendages to successfully orient. Animals using information from the bilaterally paired appendages in the control group exhibited increased walking speed, increased speed to source and decreased heading angles towards the source compared to these measurements taken from lesioned groups. There was no significant difference in any parameters between animals with unilateral or bilateral lesions. This strongly suggests that these animals are reliant on the spatial comparison of differences between bilaterally paired olfactory appendages for successful orientation.     

Preemptive legislation inhibits the anthropogenic spread of an aquatic invasive species, the rusty crayfish (Orconectes rusticus)

The enactment of legal policies is often recommended to prevent anthropogenic introductions of invasive species. In this paper, we evaluated the effectiveness of proactive state legislative policies in deterring colonization by rusty crayfish (Orconectes rusticus) and the expected spread into previously uninvaded states using network-based spatial analysis. We found that the presence of regulations was positively associated with the presence of rusty crayfish (p = 0.027), but often regulations were put into effect subsequent to the invasion. Regulations that did not explicitly prohibit transport, applied to specific drainages, or prohibited only rusty crayfish were not effective. However, preemptive legislation was effective in reducing the likelihood of invasion, if regulations prohibited the transport of all live crayfish species between water bodies, as only 1 state which passed such legislation prior to invasion by rusty crayfish was subsequently colonized (S = 12, p = 0.031). Five states are likely to be invaded by rusty crayfish via range expansion across interstate drainages, and dispersal rates suggest that all of these states could be colonized within 10 years. While it is unlikely that regulations will prevent dispersal-based invasion across state lines through shared drainages, preemptive legislation can significantly reduce anthropogenic spread of aquatic invasive species between drainages and effectively retard the expansion of rusty crayfish. Our results suggest that the most effective form of legislation is one that does not require individuals to identify species, thus we recommend states enact policies that explicitly prohibit the transport of all live crayfish between water bodies.     

The effect of aquatic plant chemistry and morphology on feeding selectivity by the crayfish, Orconectes virilis

SUMMARY. 1. Feeding preference trials conducted with male and female crayfish ( Orconectes virilis ) showed that out of ten submersed aquatic plant species, Chara sp. and Lemna trisulca were the preferred food sources, followed by Myriophyllum exalbescens or Utricularia vulgaris .
2. Analysis of plant fibre, alkaloid, total phosphorus, total and available nitrogen, and organic content showed that crayfish feeding preferences were not related ( P >0.5) to plant fibre or alkaloid content but were negatively correlated ( P <0.05) with plant nutrient content such that species of low nutritional value ( Chara and Lemna ) were consistently preferred.
3. Despite this preference for low nutrient food, daily intake rates by crayfish for phosphorus, nitrogen and organic matter were greatest when feeding on Lemna or Chara due to the large quantity consumed.
4. We suggest that feeding preferences of O. virilis are not determined by plant chemistry but, rather, relate to ease of handling small, short, bottom-dwelling plants such as Chara and Lemna as compared with large erect, roseate or floating-leaved forms. Because crayfish are selective herbivores, their introduction into waters where they do not naturally occur could have a dramatic effect on both the species composition and biomass of submersed plants.     

Invasion of an occupied niche by the crayfish Orconectes rusticus: potential importance of growth and mortality

We are exploring mechanisms of an invasion that contradicts the oft-cited generalization that species invade vacant niches. In northern Wisconsin lakes, the introduced crayfish Orconectes rusticus is replacing two ecologically similar resident congeners, O. virilis and O. propinquus. In laboratory experiments, we compared growth and mortality of individually maintained crayfish offered one of five ad libitum diets: invertebrates, macrophytes, dentritus, periphyton or all items combined. Mortality was highest for O. virilis and lowest for O. rusticus. Macrophyte diets yielded the highest mortality. All three species grew best on invertebrate and combination diets but grew little or not at all on diets of periphyton, detritus or macrophytes. O. rusticus and O. virilis grew more than O. propinquus. O. rusticus grew more quickly and/or was better able to survive overall than its congeners. Therefore, O. rusticus would probably have advantages over O. virilis and O. propinquus in competitive interactions, reproductive success and avoiding size-selective fish predation. Subtle interspecific differences may interact strongly with other ecological factors and contribute to the displacement of resident species from a well-occupied niche.     

Effects of declining calcium availability on the survival,growth and calcium content of a freshwater crayfish,Orconectes virilis

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