Homeostatic responses in the gut of Porcellio scaber (Isopoda: Oniscidea) optimize litter degradation

In order to examine the influence of differences in food conditions on gut characteristics in Porcellio scaber, pH-manipulated and microbially inoculated leaf litter from three different tree species were offered. Microbial activity was clearly influenced by the pH levels of the leaves. Analyses of the pH levels in the gut indicated the ability of P. scaber to buffer the pH value in the intestinal tract to about 5.5–6.0 in the anterior hindgut, and to about 6.0–6.5 in the posterior hindgut. The pH levels of the gut sections remained in this range, within a range of food pH from 4.0 to 7.5, no matter what kind of leaves the animals were fed. Homeostatic responses to changes in food pH guarantee optimized digestion of leaf litter. However, when the pH level of the litter dropped below 3.5, P. scaber was not able to maintain the pH conditions in the gut. Furthermore, microorganisms colonizing the litter biased the pH level in the anterior hindgut where digestive processes mainly take place. These results indicate a decline of litter quality with regard to the nutrition of terrestrial isopods, caused by acidification and consequently reduced microbial activity. Accepted: 19 July 1997     

Exoskeletal cuticle differentiation during intramarsupial development of Porcellio scaber (Crustacea: Isopoda)

Exoskeletal crustacean cuticle is a calcified apical extracellular matrix of epidermal cells, illustrating the chitin-based organic scaffold for biomineralization. Studies of cuticle formation during molting reveal significant dynamics and complexity of the assembly processes, while cuticle formation during embryogenesis is poorly investigated. This study reveals in the terrestrial isopod Porcellio scaber, the ultrastructural organization of the differentiating precuticular matrices and exoskeletal cuticles during embryonic and larval intramarsupial development. The composition of the epidermal matrices was obtained by WGA lectin labelling and EDXS analysis. At least two precuticular matrices, consisting of loosely arranged material with overlying electron dense lamina, are secreted by the epidermis in the mid-stage embryo. The prehatching embryo is the earliest developmental stage with a cuticular matrix consisting of an epicuticle and a procuticle, displaying WGA binding and forming cuticular scales. In newly hatched marsupial larva manca, a new cuticle is formed and calcium sequestration in the cuticle is evident. Progression of larval development leads to the cuticle thickening, structural differentiation of cuticular layers and prominent cuticle calcification. Morphological characteristics of exoskeleton renewal in marsupial manca are described. Elaborated cuticle in marsupial larvae indicates the importance of the exoskeleton in protection and support of the larval body in the marsupium and during the release of larvae in the external environment.     

Ultrastructural study of yolk formation in Porcellio scaber Latr. (Isopoda)

Two types of yolk are formed in the developing oocytes of a terrestrial crustacean, Porcellio scaber. The intra-oocytic yolk, arises through autosynthesis, and the extra-oocytic yolk, is derived from micropinocytosis. the so-called disc shaped bodies, which occur in large numbers within the cisternae of a branched system of endoplasmic reticulum, are precursors of the intra-oocytic uolk. Dictyosomes are not involved in yolk formation in this species. Thus, the vitellogenesis of Porcellio scaber occurs in a manner analogues to that described in the aquatic crustaceans, which indicates that environmental factors have relatively little effect on this process.     

Microscopic anatomy and ultrastructure of the stomach of Porcellio scaber (Crustacea,Isopoda)

Summary From the micromorphological viewpoint, the stomach is by far the most complicated part of the digestive tract of Porcellio scaber. All parts are shown in electron micrographs. The simplest inner surface of the stomach is the dorsal part, which has an unpaired flap directed posteriorly (lamella dorsalis). The lateral walls carry the lateralia, superolateralia and lamellae annulares. The most complicated region of the stomach is the ventral surface with its anterior primary filter, the adjacent masticatory areas, the posterior secondary filter on the lateral sides of the inferomedianum and the inferolateralia. Ingested food can be filtered twice: first on the primary filter and then on the secondary filter. The digestive fluid secreted in the midgut glands can be conveyed on the same route as the filtrate of the gastric juice, but in the other direction. The posterior part of the stomach is wrapped by a ring-shaped fold of the anterior hindgut, thus making seven epithelia closely apposed to each other.     

Personality affects defensive behaviour of Porcellio scaber (Isopoda,Oniscidea)

We evaluated individual behavioural patterns of isopods expressed as tonic immobility following some intrusive treatments. Common rough woodlice, Porcellio scaber, were kept individually in plastic boxes and tested for tonic immobility repeatedly. Reactivity, sensitivity (number of stimuli needed to respond), and endurance of tonic immobility (TI) according three types of treatments (touch, squeeze, drop) were evaluated. Touch was the weakest treatment and it was necessary to repeat it a number of times to obtain a response; while squeeze and drop induced TI more frequently. Nevertheless, duration of the response persisted for a longer time with the touch treatment. Within each set of the three treatment, the strongest response was the third one, regardless of treatment type. Duration of reaction was affected by the size of the woodlouse, the smallest individuals feigning death for the shortest time. Despite body size, we found a significant individual pattern of endurance of TI among tested woodlice, which was stable across treatments as well as across time (5 repetitions during a 3 week period). Porcellio scaber is one of the first species of terrestrial isopods with documented personality traits.     

Metallothionein-like proteins and zinc-copper interaction in the hindgut of Porcellio scaber (crustacea: Isopoda) exposed to zinc

Metallothioneins (MTs) are ubiquitous low-molecular-weight metalbinding proteins, with a variety of functions in metal metabolism ascribed to them. Among terrestrial invertebrates, MTs have been studied in nematodes, insects, snails, and earthworms. The aim of this study was the characterization of MT-like proteins in the terrestrial isopod crustacean Porcellio scaber in order to analyze their probable role in the metaboliss of copper (Cu) and zinc (Zn). Dietary Zn supplementation (793 μg Zn/g dry food, 6 d) was applied to stimulate MT synthesis. After separation of the hindgut postmicrosomic supernatant (cytosol) of Zn-exposed animals by gel filtration on a Sephadex G-75 column, a Cu- and Zn-containing peak was detected in the position of V _c/V_o≈2, where MTs are expected to elute. Rechromatography of these fractions by size-exclusion chromatography-high-performance liquid chromatography revealed that the 215-nm absorbance peak coincided with the absorbance peak of the rabbit MT II standard. These low-molecular-weight Cu- and Zn-binding compounds, detected in the cytosol of the hindgut cells in Zn-exposed P. scaber. are considered to be Cu, Zn-MT-like proteins. To our knowledge, this is the first report on the characterization of MT-like proteins in isopod crustaceans. These results also indicate that both Zn and Cu dynamics in P. scaber hindgut are affected at the given dietary Zn supplementation and that MT-like proteins are involved in this Zn-Cu interaction.     

Shelter site use and secondary sex ratios in the woodlice Armadillidium vulgare and Porcellio scaber (Crustacea: Isopoda)

In grassland populations of the woodlice Armadillidium vulgare and Porcellio scaber (Latreille) the sex ratio in hand collections from beneath wooden offcuts and roofing tiles varied through the breeding season. The proportion of males was higher in these collections from surface refugia than in independent estimates of the population sex ratio taken from soil cores. Collections became female-biased in June and July when most females were carrying broods, most notably in P. scaber . It is suggested that males use surface shelter between foraging events. A. vulgare males are particularly abundant under boards in May as they become active on the soil surface in order to locate mates. Females may find the higher and more equable temperatures in refugia attractive because they reduce brood development time but the cost of this advantage is the risk of failure to balance moisture requirements. These patterns of habitat use by the sexes have important implications for several population processes, the evolution of life-history tactics and the development of mating systems in these species.     

Epithelial thickness and lipid droplets in the hepatopancreas of Porcellio scaber (Crustacea: Isopoda) in different physiological conditions

We investigated the morphometric characteristics of the hepatopancreatic epithelium in the terrestrial isopod Porcellio scaber during acclimatization to laboratory conditions, during the daily cycle, the molt cycle, and fasting. The hepatopancreatic epithelium was analyzed using computer-assisted microscopy of serial sections of the hepatopancreatic tubes. In addition, the abundance, the distribution, and the size of lipid droplets in the hepatopancreatic epithelium were recorded. The experimental animals were collected in the field and transferred to the laboratory. The hepatopancreatic epithelium was thinner and lipid droplets reduced after 2 months of acclimatization to laboratory conditions. The daily cycle and the molt cycle affected neither the epithelial thickness nor the abundance of lipid droplets. But in animals fasted for 2 weeks, these two parameters were significantly reduced. Based on both the epithelial thickness and the abundance of lipid droplets in B cells, we propose criteria for estimating the stress status of the animals. With the possibility to determine the stress status, many studies on isopods gain in relevance.     

Interaction of lead and calcium uptake by the woodlouse,Porcellio scaber (Isopoda,Porcellionidae)

Summary Lead and calcium nitrate in a potato base were fed to Porcellio scaber; three levels of each cation were used in a 3² factorial design experiment. The amounts accumulated were determined by atomic absorption spectrophotometric analysis of nitric acid digests of whole animals. Both cations increased markedly in the woodlice with increasing dosage and a clear correlation was found between the rates of uptake of the two.     

Ultrastructure of the digestive system and the fate of midgut during embryonic development in Porcellio scaber (Crustacea: Isopoda)

Microscopic anatomy of the digestive system in embryos and larvae of the terrestrial isopod crustacean Porcellio scaber was investigated by light bright field, fluorescence and electron microscopy. During marsupial ontogenetic development the event-dependent staging was used to discriminate the various embryonic stages. At the late embryo stage the differentiation of the ectodermal part of the gut into the complex filtering foregut and the hindgut with absorptive and transporting functions is accomplished. The gut of the marsupial manca larva is fully developed and similar to that of the adult. In early embryos the endodermal midgut gland primordia are filled with yolk and lipid globules. In late embryos the epithelium of paired midgut gland tubes is composed of two cell types; one of them exhibits orange autofluorescence. The endodermal cells located between the foregut and the midgut glands of late embryos form the prospective midgut. The cells have electron dense cytoplasm, abundant glycogen fields, endoplasmic reticulum, dictyosomes and numerous vesicles. In the adults the endodermal cells of the midgut remain only in the midgut gland ducts which connect the midgut glands and the foregut. Details of the cellular ultrastructure and morphogenesis of the ectodermal and endodermal parts of the digestive system during embryonic development of Porcellio scaber provide data for further phylogenetic and comparative studies in peracaridan crustaceans and other arthropods.     

Architecture of the organic matrix in the sternal CaCO3 deposits of Porcellio scaber (Crustacea, Isopoda)

Before the molt terrestrial isopods resorb calcium from the posterior cuticle and store it in large deposits within the first four anterior sternites. In Porcellio scaber the deposits consist of three structurally distinct layers consisting of amorphous CaCO3 (ACC) and an organic matrix that consists of concentric and radial elements. It is thought that the organic matrix plays a role in the structural organization of deposits and in the stabilization of ACC, which is unstable in vitro. In this paper, we present a thorough analysis of the ultrastructure of the organic matrix in the CaCO3 deposits using high-resolution field-emission scanning electron microscopy. The spherules and the homogeneous layer contain an elaborate organic matrix with similar structural organization consisting of concentric reticules and radial strands. The decalcification experiments reveal an inhomogeneous solubility of ACC within the spherules probably caused by variations in the stabilizing properties of matrix components. The transition between the three layers can be explained by changes in the number of spherule nucleation sites.     

Morphology,morphometry and ultrastructure of the maxillary gland of the terrestrial isopod Porcellio scaber (Crustacea,Oniscidea)

Summary The ultrastructure of the maxillary gland of the terrestrial isopod Porcellio scaber is described. The gland is composed of an end sac, an excretory duct and a terminal duct which opens by a valve at the base of maxilla 2. An epithelium of podocytes in the end sac enables passive ultrafiltration by haemolymph pressure. The excretory duct shows ultrastructural adaptations to secretion and resorption. SEM micrographs reveal the location and the morphology of the valve at the excretory pore. A model reconstructed from serial sections allowed the calculation of morphometric data of the gland. The ultrafiltration area of the end sac and the area of resorption and secretion of the excretory duct amount to 0.091 and 0.157 cm² per 1 g of fresh body weight, respectively. The total volume of the gland is calculated to be 0.04 mm³ in a specimen of 13.7 mm length. In comparison with the marine species Mesidotea entomon, the relative areas for ultrafiltration and resorption of the gland of P. scaber are more than twice as large as the corresponding areas of the marine species. This relative enlargement of the gland in P. scaber and the form of the valve at the excretory pore are seen as adaptations to terrestrial life.     

Amorphous and crystalline calcium carbonate distribution in the tergite cuticle of moulting Porcellio scaber (Isopoda, Crustacea)

The main mineral components of the isopod cuticle consists of crystalline magnesium calcite and amorphous calcium carbonate. During moulting isopods moult first the posterior and then the anterior half of the body. In terrestrial species calcium carbonate is subject to resorption, storage and recycling in order to retain significant fractions of the mineral during the moulting cycle. We used synchrotron X-ray powder diffraction, elemental analysis and Raman spectroscopy to quantify the ACC/calcite ratio, the mineral phase distribution and the composition within the anterior and posterior tergite cuticle during eight different stages of the moulting cycle of Porcellio scaber. The results show that most of the amorphous calcium carbonate (ACC) is resorbed from the cuticle, whereas calcite remains in the old cuticle and is shed during moulting. During premoult resorption of ACC from the posterior cuticle is accompanied by an increase within the anterior tergites, and mineralization of the new posterior cuticle by resorption of mineral from the anterior cuticle. This suggests that one reason for using ACC in cuticle mineralization is to facilitate resorption and recycling of cuticular calcium carbonate. Furthermore we show that ACC precedes the formation of calcite in distal layers of the tergite cuticle.     

The postmarsupial development of Porcellio siculoccidentalis, with some data on reproductive biology (Crustacea, Isopoda, Oniscidea)

In the broader context of research on the Sicilian Porcellio imbutus-complex, the postmarsupial development of Porcellio siculoccidentalis Viglianisi, Lombardo & Caruso, 1992 was studied in detail. This research was conducted in the laboratory under controlled conditions, allowing us to follow the stages of development, from the formation of the marsupium in ovigerous females until the larval stages and development of the seventh pair of legs. The timing of developmental stages and the morphological modifications of appendages in the postmarsupial manca stages (M I-M III) are described. The manca stage M I had a duration of about one hour. Ovigerous females were collected and reared separately, and the number of parturial molts in the absence of males was counted. The results showed a maximum of four successive parturial molts. Fecundity and fertility were evaluated as the number of eggs and embryos, respectively, inside the marsupium of the ovigerous females. Both parameters were positively correlated with the size of the females. The maximum numbers of eggs and embryos in the marsupium were 113 and 141, respectively. Data describing the total number of postmarsupial mancas released per month indicated that the highest release occurred in April.     

Population dynamics and reproductive aspects of Porcellio albinus (Isopoda,Oniscidea) of Zarat (Gabes,Tunisia)

Porcellio albinus is a nocturnal species inhabiting xeric habitats in Tunisia. Size distribution, population dynamics, breeding activity, fecundity, fertility, reproductive allocation (RA), and life span were studied to establish the reproductive strategy of a population of this species in the sandy coastal area of Zarat. Monthly sampling was undertaken between July 2012 and November 2013. The overall sex-ratio of P. albinus was female biased. The relative abundance of adults, much more representative than that of juveniles, might be the consequence of the absence of specimens less than 8 mm in size in all samples collected outside the burrows at night. Reproduction is seasonal in P. albinus with two breeding seasons, the first one from March to June and the second in September. Five cohorts were recruited per year, four during the spring and one during the fall. The number of eggs per female ranged from 13 to 67 and was positively correlated with female body size. Fertility varied from 5 to 39 mancae. Compared to other xeric species, RA was low to produce a brood, but high to release a single manca. The results provide a better understanding of the reproductive behavior of this terrestrial isopod, living under severe environmental conditions.     

Microplastic ingestion affects hydrogen production and microbiomes in the gut of the terrestrial isopod Porcellio scaber

Microplastic (MP) is an environmental burden and enters food webs via ingestion by macrofauna, including isopods (Porcellio scaber) in terrestrial ecosystems. Isopods represent ubiquitously abundant, ecologically important detritivores. However, MP-polymer specific effects on the host and its gut microbiota are unknown. We tested the hypothesis that biodegradable (polylactic acid [PLA]) and non-biodegradable (polyethylene terephthalate [PET]; polystyrene [PS]) MPs have contrasting effects on P. scaber mediated by changes of the gut microbiota. The isopod fitness after an 8-week MP-exposure was generally unaffected, although the isopods showed avoidance behaviour to PS-food. MP-polymer specific effects on gut microbes were detected, including a stimulation of microbial activity by PLA compared with MP-free controls. PLA stimulated hydrogen emission from isopod guts, while PET and PS were inhibitory. We roughly estimated 10⁷ kg year⁻¹ hydrogen emitted from the isopods globally and identified their guts as anoxic, significant mobile sources of reductant for soil microbes despite the absence of classical obligate anaerobes, likely due to Enterobacteriaceae-related fermentation activities that were stimulated by lactate generated during PLA-degradation. The findings suggest negative effects of PET and PS on gut fermentation, modulation of important isopod hydrogen emissions by MP pollution and the potential of MP to affect terrestrial food webs.