The influence of intrinsic and extrinsic factors on developmental parameters and their relationships in the marine isopod Idotea linearis (Crustacea)

Developmental and reproductive parameters and their relationships were studied in the marine isopod Idotea linearis. We hypothesized that (1) the temporal patterns of molting and growth undergo complex and sex-specific changes with age as well as with the onset of sexual maturation, and that (2) sexual maturation (and dependent parameters) is controlled by the photoperiod. Both males and females were singly cultured in the laboratory at two alternative photoperiods (constant long and short days, respectively) from hatching until death. Males molted and grew throughout their life, showing a steady increase in stage duration and body size with each molt. Females, in contrast, showed much more complex modifications in molt chronology due to reproductive demands. There was some variability in the stage number, when females reached maturity. Reaching maturity early in the succession of molts was associated with smaller body size at maturity, smaller size of broods, but higher average number of broods per lifetime. Post-puberty molts in females occurred without further growth, and successive broods did not differ in size. The photoperiod strongly affected sexual maturation (and thus in turn molting and growth patterns) in females, while males remained completely unaffected by the photo regime.     

Mechanics of branchial ventilation in the valviferan isopod Idotea wosnesenskii (Crustacea)

Idotea wosnesenskii Brandt, a valviferan isopod, has five pairs of pleopods within a branchial chamber enclosed by modified uropods (operculae). The pumping mechanism of resting isopods was videotaped to observe ventilation, and particularly to determine whether or not the swimming pleopods (first three pairs) and the gas exchange pleopods (last two pairs) have ventilatory roles. Observations on intact animals (to observe water flow into and out of the branchial chamber) and on animals with operculae propped open or with parts of their abdominal wall removed (to study the actual pumping movements of the pleopods) revealed that all five pairs of pleopods function in ventilation. The ventilatory stroke has two phases: an opening phase in which the pleopods move medially and ventrally, opening spaces between pleopods (analogous to the swimming recovery stroke), and a closing phase, wherein the pleopods move laterally and dorsally, reducing the gaps between succeeding pleopods (analogous to the swimming power stroke). Ventilatory strokes may be continuous or they may be separated by resting phases. Both the frequency and amplitude are variable: the frequency changes greatly due to the wide variation in resting phase duration. By opening the operculae the amplitude can increase so that ventilatory strokes grade into swimming strokes. The ventilatory stroke pattern at rest is generally similar to the swimming stroke pattern but differs in its slower and nearly simultaneous dorsoventral movements and its lower frequencies. Dye and particle movements around intact animals and those with exposed pleopods show that water typically takes about three strokes to pass through the branchial chamber.     

Characterization of polymorphic microsatellite loci in the marine isopod Sphaeroma terebrans (Crustacea, Isopoda)

Nine polymorphic microsatellites were characterized in the marine isopod Sphaeroma terebrans (Isopoda, Sphaeromatidae) for phylogeographic and parentage analyses. The number of alleles ranged from 3 to 10. Observed heterozygosity ranged from 0.428 to 0.950, while expected heterozygosity ranged from 0.532 to 0.889. Heterozygote deficiency was detected for one locus, possibly the result of null alleles.     

Differential predation on sexes affects colour polymorphism of the isopod Idotea baltica (Pallas)

Variable selection, including spatio-temporal variation, frequency-dependent selection and differential selection due to habitat choice, may maintain polymorphism in heterogeneous environments. We studied predation as a selective agent on colour polymorphism of the aquatic isopod I baltica. Variable predation on this species can arise from at least three sources. First, apostatic selection was studied by testing the formation of preferences on colour morphs in the perch, a common predator of I baltica. Such acquired preferences should induce apostatic selection. While our results indicate some acquired preferences, there was significant heterogeneity in the behaviour of predator individuals. Second, temporal variation in selection can arise due to habitat shift from the green algae juvenile habitat to the bladderwrack adult habitat, and the consequent change in the crypsis of the morphs. Different crypsis between sexes probably promoted high predation mortality among females in the juvenile habitat. The high rate of male mortality during the breeding period, on the other hand, was presumably due to their high mate-searching activity. Third, the sex-dependent habitat choice of I baltica leads to sexual differences in the susceptibility of morphs to predation. Predators preferred the white-spotted morph over the uniform one in males but not in females, supporting the 'dimorphic niche' hypothesis as an explanation of sexual differences in morph frequencies. Finally, no evidence was found that the colouration patterns were under sexual selection. We therefore conclude diat variable predation is the most promising explanation for the maintenance of polymorphism in I. baltica.     

Reckless males, rational females: dynamic trade-off between food and shelter in the marine isopod Idotea balthica

Habitat choice of herbivores is expected to be a resolution of a trade-off between food and shelter. The resolution of this trade-off may, however, be dynamic within a species because distinct phenotypes may value these factors differently and the value may vary temporally. We studied this hypothesis in the marine herbivore Idotea balthica (Isopoda), by simultaneously manipulating both food and shelter, and investigated whether the resolution of the trade-off differed between sexes, colour morphs and day and night (i.e. high and low predation risk). Isopods chose between exposing and concealing backgrounds in which the quantity or quality of food varied. When choosing between the backgrounds in the absence of food, females preferred the concealment more than males did. However, in a trade-off situation the isopods traded shelter for food, and females more so than males. Thus, males' lower preference for the shelter was not counterbalanced by a stronger preference for food. The microhabitat use also differed between night and day showing adaptation to diurnally fluctuating predation risk. We suggest that microhabitat utilization of females is more strongly tied to variation in risk and resources than that of males, for whom other factors, such as seeking mates, may be more important.     

Respective influence of extrinsic and intrinsic factors on the age-related decrease of thymic secretion.

The serum level of a circulating thymic factor (FTS) described in our laboratory diminishes in mice after adult thymectomy and with age. However, thymuses from old mice, when grafted into young adult thymectomized recipients lacking circulating FTS, can still partially restore the circulating FTS level of the recipients, whereas newborn thymuses are less efficient in restoring the serum level of FTS in old recipients than in young adult thymectomized recipients. Taken together, our results suggests that in addition to an intrinsic deficiency of the thymic secretion, "environmental" factors play a role in the disappearance of circulating FTS with age, the more so since we observed in old mouse sera factors inhibiting the in vitro biologic activity of FTS, factors that are absent from young mouse sera.     

Burrow plasticity in the deep-sea isopod Bathynomus doederleini (Crustacea: Isopoda: Cirolanidae)

We investigated whether the deep-sea isopod Bathynomus doederleini has the capacity to change burrow length in response to changes in environmental conditions. We observed burrowing behavior in individuals that were placed on substrates with either simple (ST) or complex (CT) surface topographies. Individuals in the ST group (N = 10) constructed seven burrows. The mean ratio of the burrow length to body length was 1.8. The individuals in the CT group (N = 10) constructed eight burrows with a mean ratio of burrow length to body length of 2.5. Thus the burrows were significantly longer in the CT group. In addition, the isopods in the CT group often incorporated a chamber in the mid-section of the burrow. Our results may be used to infer the determinants of burrow morphology and speculate about the lifestyle of this species in the deep sea.     

Male dimorphism in the harem-forming gnathiid isopod Elaphognathia discolor (Crustacea: Isopoda)

Previously unreported males of a gnathiid isopod were found in reproductive aggregations of the harem-forming gnathiid Elaphognathia discolor. Although the male gnathiids were small in size and morphologically different from E. discolor males, the male sexual organ, appendix masculina, was similar to that of E. discolor males, and possible conspecific larvae and females of the small male gnathiid were never found. In the laboratory, the small male gnathiids as well as male E. discolor successfully copulated with female E. discolor, and the development of embryos in female brood pouches was observed. Offspring of small male gnathiids develop to adults of E. discolor after molting three times, or small male gnathiids after molting two times. Thus, the small male gnathiid was concluded to be an alternative male form compared to the regular large male form of E. discolor. This male polymorphism was thought to have a genetic basis, since no small male specimens appeared in offspring of regular E. discolor males. Field sampling showed that a regular large male formed a harem composed of one large male and several females and never coexisted with other large males as previously reported. However, small males were often found together with large males. Therefore, small males are thought to be sneakers intruding into harems dominated by large males.     

Evolutionary biogeography of the Microparasellid isopod Microcharon (Crustacea) in the Mediterranean Basin

The interstitial subterranean isopod Microcharon (Crustacea) is highly diversified within the Mediterranean. The present distribution of 70 species is considered the result of the joint history of this genus and its environment. This stygobiont is derived from marine surface ancestors. Both Plate Tectonics and the two-step model of colonization and evolution, the second phase of which represents vicariance during the Tethys regressions, enable an understanding of the evolutionary history of this monophyletic group. The single extant marine littoral species was separated from its sister group by the closure of the Gibraltar Straits during the Messinian. The drift of the Apulian plate, the Alboran-Kabylian-Calabrian fragment as well as the opening of the western Mediterranean through the translation of the Corsican-Sardinia plate could explain the divergences in the western and the eastern Mediterranean. The primitive sister groups in Morocco and in Spain would have been left by the Turonian Tethys regression. Such regressions of the Tethys embayments could have played a major role in the evolution of the genus from Turanian up to Tortonian in Norht Africa, and up to Pliocene and even to Pleistocene in Italian, Iberian and Greek zones. Several eastern European species may be a product of the Paratethys regressions. Everywhere, species with plesiomorphic characters are related to old regressions and vice versa. Local geological events could have resulted in further vicariance divergences. There is a good congruence between the evolutionary history of Microcharon and other stygobiont isopod, amphipods or syncarids living in the same regions studied in the Mediterranean.Presented at the Symposium of Biogeography at Subterranean Crustaceans: the effects of different scales (June 1992; Charleston, South Carolina, U.S.A.). Manuscrips prepared for print by J. R. Holsinger.     

Product conformation driven splicing of unprotected peptides by reverse proteolysis: influence of intrinsic and extrinsic factors

The structural motif of 'product conformation driven V8 protease catalyzed ligation reaction' can be represented by FR(I)-EALER-FR(II). The relative roles of the flanking regions (FR(I) and FR(II)) and of splicedon, the central penta-peptide, on the thermodynamic stability of the 'conformational trap' of the product has been now evaluated as a function of co-solvent concentration. The studies have established that the thermodynamic stability of the conformational trap of alpha17-40des23-26 with four different splicedons (EALER, EALEV, EYGER, or EGAER) that differ in the intrinsic alpha-helical potential of their amino acid residues and/or ability to generate i, i+4 side chain interaction is a direct correlate of the n-propanol induced alpha-helical conformation of the product. On the other hand, when the product is defined by only splicedon EALER, and the flanking regions are disitinct; no correlation could be drawn between the stability of the trap and solvent induced alpha-helical conformation, even though these generally give an equilibrium yield of 45% in 30% n-propanol and is not influenced by an increased propanol concentration. However, when the splicedon EALER with given FR(I) and FR(II) region develops a 'conformational trap' of a lower stability in 30% propanol as seen with beta18-25(A22)-EALER-beta31-39, the stability increases in 60% n-propanol, without significant increase in the alpha- helical conformation. Though, primary structure of RNAse1-20, could be presented as RNAse1-5-AKFER- RNAse1-20, and alpha-helical conformation is induced to this peptide both in 30 and 60% propanol, splicedon AKFER by itself does not develop the 'conformational trap' of RNAse1-20. The splicedon AKFER of RNAse1-20 fails to develop the 'conformational trap', due to an intrinsic inhibitory potential of its FR region, RNAse11-20; replacing RNAse11-20 with alpha32-40 enables the splicedon AFKER to generate the 'conformational trap'. The studies presented here have demonstrated the primary role of flanking regions in establishing the amount of the solvent induced alpha-helical conformation and that of the splicedon in dictating the thermodynamic stability of its 'conformational trap' of the products, nonetheless one influences the other to some degree. We suggest that the stability of the 'conformational trap' of the product reflects the ability of the splicedon to 'recruit' the product conformation to protect the spliced peptide bond, i.e. to reduce the helix-coil transition of the spliced region which in turn imparts a degree of resistance to the spliced peptide bond.     

Natural history of the Pan-American sand beach isopod Excirolana braziliensis (Crustacea: Malacostraca)

A population of the intertidal isopod Excirolana braziliensis was followed over a 10-month period at Naos Island, Panama. Average intertidal density was 2147/m² with maximum abundance occurring in the highest level. Adults were present only in the highest tidal level. Juveniles were distributed throughout all tidal levels and comprised at least 70% of the population at all times. Mean size of individuals decreased from the highest level seaward. Reproductive females were present the year round. The number of young ranged from four to 17 per brood and was dependent on the length of the female. Adults were very tolerant of changes in temperature and salinity.     

Ultrastructure of the pleopods of the estuarine isopod Cyathura carinata (Crustacea: Isopoda: Anthuridea)

Summary The exopodal branches of the pleopods of Cyathura carinata have a thicker cuticle than the endopodites, but the hypodermis proved to be very thin and probably of respiratory function. The nuclei are found in pilaster areas together with stabilizing elements (micro-tubuli attached to cuticular pegs), which are typical for hypodermal tendon cells of other arthropods. Precipitation of Ag₂S revealed that the cuticle of the exopod is more permeable to silver ions than the cuticle of the endopods. The endopodal epithelia have the features of typical osmoregulating tissues (apical folds, mitochondrial pumps, and basal invaginations of the plasma membrane) and reduce silver nitrate. The stabilizing pilaster areas are composed of the same elements as in the exopods.     

Mathematical modelling of residue allelopathy: the effects of intrinsic and extrinsic factors

The mechanistic model of residue allelopathy that was constructed before only simulates allelopathic phenomena caused by decaying plant residues and does not include factors controlling expression of such phenomena. As both intrinsic and extrinsic factors are affecting the potential of residue allelopathy, the model has been developed further to include these factors. The concept of a constant for residue-allelopathic potential, functions for effects of temperature, water, and their combination on allelochemical concentrations were added to the model. As a result, the improved residue model is able to examine the responses to these phenomena under a wide variety of conditions. Several sets of published experimental data were compared with prediction from the model, and a good agreement has been achieved. Its significance in helping better understanding of residue allelopathy is discussed.     

Effect of intrinsic and extrinsic factors on the variability of echolocation pulses of Myotis nigricans (Schinz, 1821) (Chiroptera: Vespertilionidae)

Bats may exhibit plasticity in echolocation pulses as response to intrinsic and extrinsic factors, and the estimation of the magnitude of such variation can provide confidence in acoustic monitoring. Myotis nigricans (Schinz, 1821) is a widely distributed but relatively understudied Neotropical species for which, during fieldwork, we found maternity colonies in the Lagunas de Montebello National Park, in Chiapas, South-east Mexico, and no previous information in the area. Therefore, we aimed to provide an acoustic characterization on the basis of intraspecific variability for its recognition using bat detectors. For this purpose, we examined the moulding of shape, frequency-based and time-based acoustic parameters, specifically by the effect of age group (sub-adults vs. adults) and acoustic environment (open space vs. background vegetation). By graphic comparison of echolocation pulses between acoustic environments, we observed changes in shape by an increase in bandwidth and steeper modulation along background vegetation. Statistically, on univariate basis, we did not find a significant effect of age group, but we did of acoustic environment, specifically on highest frequency (higher average), duration (shorter average) and interpulse interval (shorter average) along background vegetation. On multivariate basis, we confirmed shorter average interpulse interval along background vegetation. The overall classification accuracy was relatively high (82.22%): 80% in open space and 84% along background vegetation. Our work reinforces previous knowledge about sound constraints imposed by vegetation clutter, and provides a reliable framework for acoustic monitoring of this species across structurally variable, and hence acoustically variable, environments in the area.     

Anatomy of the extrinsic gut musculature of Gammarus minus (Crustacea: Amphipoda)

Inserting on the buccal and esophageal foregut of Gammarus minus are numerous pairs of serially arranged dorsal dilator muscles, a single pair of lateral muscles, and two pairs of posterior muscles. Muscles of the cardiac stomach include three dorsal sets, a single pair associated with the pterocardiac ossicles, and two pairs inserting on the ventral aspect. A single pair of muscles inserts on the lateral aspect of the pyloric stomach. The extrinsic muscles of the foregut originate from exoskeletal apodemes of the cephalothoracic cuticle, sockets of the mandible, and a maxillary bridge that lies just ventral to the cardiac stomach. The extrinsic musculature of the hindgut is restricted to the rectal region and consists of paired dorsal and ventral series in an X-configuration. A single unpaired muscle inserts on the ventral midline. Extrinsic muscles of the hindgut originate from the integument of the last pleonic segment. The general arrangement of extrinsic gut muscles in G. minus is similar but not identical to that of other amphipods studied. However, the pattern is quite different from that of other malacostracans.     

Observations with a different approach to morphological colour variation in an isopod population (Crustacea: Isopoda)

Morphological colour variation in Idotea baltica basteri Audouin, 1827 (Isopoda: Crustacea) is reported for the first time on populations from a brackish water lake (Kucukcekmece Lagoon) in Turkey. According to pigmentation characteristics, the isopods were described and sorted to ten different colour morphs which are named as maculata, maculata-lineata, uniformis, uniformis-lineata, immaculatum, immaculatum-lineata, nigrum, nigrum-lineata, albafusca, and albafusca-fasciatum. The similarity of the morphs was compared by defining digit codes to some of the visual pigmentation characteristics and calculating the Bray-Curtis similarity index of the colour morphs. The albafusca and albafusca-fasciatum forms were found to be the most similar to each other (similarity 96%), whereas immaculatum forms were considered the most different from maculata-lineata (similarity 50%). The morphs named maculata and maculata-lineata, having the highest abundance with 70%, belong to the most successful forms which may achieve crypsis by background matching in their living habitat in Kucukcekmece Lagoon which has sandy and rocky bottom surface area.