Origin and evolutionary plasticity of the gastric caecum in sea urchins (Echinodermata: Echinoidea)

Background  

The digestive tract of many metazoan invertebrates is characterized by the presence of caeca or diverticula that serve secretory and/or absorptive functions. With the development of various feeding habits, distinctive digestive organs may be present in certain taxa. This also holds true for sea urchins (Echinodermata: Echinoidea), in which a highly specialized gastric caecum can be found in members of a derived subgroup, the Irregularia (cake urchins, sea biscuits, sand dollars, heart urchins, and related forms). As such a specialized caecum has not been reported from "regular" sea urchin taxa, the aim of this study was to elucidate its evolutionary origin.     

Comparative morphology of the axial complex and interdependence of internal organ systems in sea urchins (Echinodermata: Echinoidea)

Background

The axial complex of echinoderms (Echinodermata) is composed of various primary and secondary body cavities that interact with each other. In sea urchins (Echinoidea), structural differences of the axial complex in "regular" and irregular species have been observed, but the reasons underlying these differences are not fully understood. In addition, a better knowledge of axial complex diversity could not only be useful for phylogenetic inferences, but improve also an understanding of the function of this enigmatic structure.

Results

We therefore analyzed numerous species of almost all sea urchin orders by magnetic resonance imaging, dissection, histology, and transmission electron microscopy and compared the results with findings from published studies spanning almost two centuries. These combined analyses demonstrate that the axial complex is present in all sea urchin orders and has remained structurally conserved for a long time, at least in the "regular" species. Within the Irregularia, a considerable morphological variation of the axial complex can be observed with gradual changes in topography, size, and internal architecture. These modifications are related to the growing size of the gastric caecum as well as to the rearrangement of the morphology of the digestive tract as a whole.

Conclusion

The structurally most divergent axial complex can be observed in the highly derived Atelostomata in which the reorganization of the digestive tract is most pronounced. Our findings demonstrate a structural interdependence of various internal organs, including digestive tract, mesenteries, and the axial complex.     

神农架地区4种啮齿类食性及脏器形态差异比较

生活在同一地域的物种,会因强烈的资源竞争而产生生态位重叠与分离,从而实现相互共存。以神农架地区啮齿动物为研究对象,比较分析不同啮齿类食性、脏器重量及消化道形态差异,探索其共存机制及其生理生态学上的适应策略。食性上,安氏白腹鼠(Niviventer andersoni)、社鼠(N.confucianus)、高山姬鼠(Apodemus chevrieri)、中华姬鼠(A.draco)均为杂食者,但各自喜食偏好不同,安氏白腹鼠的食物组成以植物枝叶和种子为主,社鼠、高山姬鼠和中华姬鼠的食物组成均以种子和动物性食物为主。与各自生态习性及其食性相适应,不同啮齿动物的脏器重量及消化道形态发生相应的变化。脏器重量上,不同啮齿动物心鲜重、肺鲜重、脾鲜重和干重无显著差异,其它指标差异显著。安氏白腹鼠的心干重、肺干重、肝鲜重和干重、肾鲜重和干重显著大于社鼠、高山姬鼠和中华姬鼠。消化道形态上,不同啮齿动物盲肠长、净鲜重和干重无显著差异,其它指标差异显著。其中,胃含内容物鲜重、净鲜重和干重,盲肠含内容物鲜重,大肠长、含内容物鲜重、净鲜重和干重均以安氏白腹鼠具较大值,但各自变化趋势不同;小肠长、含内容物鲜重、净鲜重和干重则以社鼠具较大值。且同属姬鼠属的高山姬鼠和中华姬鼠脏器重量及消化道形态各指标间均无显著差异。由于小泡巨鼠和猪尾鼠数量较少,未进行统计分析。以上结果显示:食物资源利用上的分化有利于安氏白腹鼠与其近缘物种社鼠及其他鼠类的同域共存,且为适应不同的食物资源利用模式,其脏器及消化道形态也发生了适应性的变化;同域分布的近缘物种高山姬鼠、中华姬鼠则有可能采取其它策略而非食物资源分化模式实现共存。     

The Digestive Tract of the Spatangoid Echinoid Echinocardium cordatum (Echinodermata): Morphofunctional Study

Abstract Echinocardium cordatum is a detritus-feeder. Its digestive tract has three functional parts: the mucus-secreting oesophagus where lubrication and sediment compaction occur, the protein-secreting stomach and gastric caecum where extracellular digestion occurs, and the absorptive non-secretory siphon, intestine, intestinal caecum and rectum. Each region harbours typical enterocytes. Acinar mucous glands occur in the posterior oesophagus prior to the incurrent aperture of the siphon; their secretions prevent sediment grains from entering the siphon and may be stimulated by epineural nerves running in the connective tissue layer. The siphon is a sucking organ, and its cytological features indicate that it is a site for absorption of dissolved organic matter. Enterocytes of the intestinal caecum show basal specializations, suggesting that it is a site of active exchange between coelom and gut. This caecum contains symbiotic sulphide-oxidizing bacteria. Collagen fibres are well developed in organs subjected to severe deformations (e.g., oesophagus and siphon); elastic-like fibres are conspicuous around haemal lacunae and at the attachment areas of gut-suspending mesenteries. Circular muscles are seen along the whole gut, whereas longitudinal muscles may be absent in areas such as the siphon. The bulky alimentary bolus that occurs in the ascending segments of the gut is transported by peristalsis; the scattered sediment grains that occur in horizontal segments are presumably moved by the contractions of the gut-suspending mesenteries.     

Microbial Distribution and Abundance in the Digestive System of Five Shipworm Species (Bivalvia: Teredinidae)

Marine bivalves of the family Teredinidae (shipworms) are voracious consumers of wood in marine environments. In several shipworm species, dense communities of intracellular bacterial endosymbionts have been observed within specialized cells (bacteriocytes) of the gills (ctenidia). These bacteria are proposed to contribute to digestion of wood by the host. While the microbes of shipworm gills have been studied extensively in several species, the abundance and distribution of microbes in the digestive system have not been adequately addressed. Here we use Fluorescence In-Situ Hybridization (FISH) and laser scanning confocal microscopy with 16S rRNA directed oligonucleotide probes targeting all domains, domains Bacteria and Archaea, and other taxonomic groups to examine the digestive microbiota of 17 specimens from 5 shipworm species (Bankia setacea, Lyrodus pedicellatus, Lyrodus massa, Lyrodus sp. and Teredo aff. triangularis). These data reveal that the caecum, a large sac-like appendage of the stomach that typically contains large quantities of wood particles and is considered the primary site of wood digestion, harbors only very sparse microbial populations. However, a significant number of bacterial cells were observed in fecal pellets within the intestines. These results suggest that due to low abundance, bacteria in the caecum may contribute little to lignocellulose degradation. In contrast, the comparatively high population density of bacteria in the intestine suggests a possible role for intestinal bacteria in the degradation of lignocellulose.     

Microanatomy of the digestive tract and accessory organs of the Japanese flathead (Inegocia japonica Cuvier, 1829) (Scorpaeniformes,Platycephalidae)

The Japanese flathead, Inegocia japonica Cuvier, 1829 is a commercially important fish in small-scale coastal fisheries in Thailand; however, an explanation of its digestive biology is missing. This study describes the digestive tract and accessory organs of I. japonica, using morphological and histological methods. The fish (10 individual fish, 24.5 ± 0.98 cm in total length) were obtained from Libong Island, Thailand. Integrated morphological and histological data showed that the digestive tract was composed of oesophagus, stomach, pyloric caeca and intestine, with accessory organs. All digestive tracts consisted of four layers, including mucosa, submucosa, muscularis and serosa. Two stomach regions were identified (cardiac and pyloric stomachs). Several clusters of gastric glands were identified in the cardiac stomach. Each gland was a unicellular structure. The apical surface of this gland contained the vacuolar cell. The intestine was lined with a simple columnar structure with goblet cells that was similar to pyloric caecum. Goblet cells were rare in the anterior intestine, in contrast to the posterior intestine where goblet cells were abundant. The numerous of hepatocyte was mostly observed in the liver, whereas an exocrine acinar cell of pancreas was also identified. The results of our observations provided the first information of the digestive tract of I. japonica and can be applied to advanced study, such as physiology and histopathology.     

禁食和重喂食对大绒鼠消化道长度与重量的影响

为探讨禁食和重喂食对栖息于横断山地区大绒鼠(Eothenomys miletus)消化道形态的影响,对禁食和重喂食条件下大绒鼠消化道各项指标进行了测定.实验分别测定了大绒鼠禁食12h、24 h、36 h和重喂食12 h、48 h、7d后的总消化道、胃、小肠、大肠、盲肠的长度及其含内容物重、去内容物重、干重.结果表明:禁食后,总消化道含内容物重、胃含内容物重和胃去内容物重增加,重喂食7d后均恢复到对照组水平.小肠含内容物重在禁食12h时最大,在重喂食12h时最小,重喂食48 h后恢复到对照组水平.禁食和重喂食条件下,大绒鼠的大肠和盲肠各指标均没有显著变化.以上结果表明,大绒鼠在食物受到限制、饥饿等胁迫因子作用下,可能通过调节消化道形态来满足部分能量需求,维持正常的生理机能.大绒鼠的消化道在禁食和重喂食中表现出的变化模式,可能与其食物资源时常波动的野外生存环境有关,从一方面反映了该物种在食物胁迫下的生存机制和适应对策.     

Fasciola gigantica: Histology of the digestive tract and the expression of cathepsin L

The digestive tract of Fasciola gigantica is composed of the oral sucker, buccal tube, pharynx, esophagus, and caecum. The tegumental-type epithelium lines the first four parts of the digestive tract while the caecal-type epithelium lines the remaining parts from the caecal bifurcation. The caecal-epithelial cells are classified into 3 types according to their staining properties and ultrastructural characteristics, as related to the amount of food contents in the caecal lumen. All caecal-type epithelial cells synthesize and secrete cathepsin L, a major group of enzymes in the digestive tract, as detected by in situ hybridization and immunolocalization. Moreover, the secreted cathepsin L is also adsorbed on the outer surface of the tegument and the glycocalyx coating of the surface of the tegument, whereas the tegumental cells and tegumental syncytium covering the parasite’s body and lining the proximal part of the digestive tract exhibit no in situ hybridization signal and immunostaining for cathepsin L.     

Evolution de quelques Enzymes Digestives chez le Jeune Mugil capito à Diverses Etapes de sa Croissance

Modifications in the digestive enzymic profile in three parts of the digestive tract (stomach, pyloric caecum and intestine) were studied during the first year of life of the teleost Mugil capito by the semi-quantitative APIZYM method. Higher enzymic activity was found int he pyloric caecum and intestine than in the stomach. Moreover, a decrease in protease activities appears during growth and is apparently due to adaptation to changes in diet.     

From mouth to anus: Functional and structural relevance of enteric neurons in the Drosophila melanogaster gut

The intestinal tract is the main organ involved in host nutritional homeostasis. Intestinal function in both vertebrates and invertebrates is partly controlled by enteric neurons that innervate the gut. Though anatomical and functional aspects of enteric neurons are relatively less characterized in Drosophila than in large insects, analyses of the role of the enteric neurons in flies have remarkably progressed in the last few years. In this review, we first provide a summary of the structure and function of the Drosophila intestine. We then discuss recent studies of the structure and function of enteric neurons in Drosophila melanogaster.     

Phospholipids of the organs and tissues of echinoderms and tunicates from peter the great bay (Sea of Japan)

The phospholipid compositions of organs and tissues were determined in representatives of two phyla of marine invertebrates: Echinodermata [Asteroidea: Aphelasterias japonica (Bell, 1881), Evasterias echinosoma Fisher, 1926, Distolasterias nipon (Doderlein, 1902), Asterias amurensis Lutken, 1871; Echinoidea: Strongylocentrotus intermedius (A. Agassiz, 1863); Holothurioidea: Cucumaria frondosa japonica (Semper, 1868), Eupentacta fraudatrix (Djakonov et Baranova, 1958), Apostichopus japonicus (Selenka, 1867)], and Tunicata [Ascidia: Halocynthia aurantium (Pallas, 1787), H. roretzi (Drasche, 1884), and Styela clava (Herdman, 1881)]. The specificity of phospholipid distribution was shown to be related to the taxonomic position of marine invertebrates and the functional properties of their organs and tissues. Ceramide aminoethylphosphonate was found only in the digestive organs of all holothurians and the starfishes D. nipon and A. amurensis, suggesting its exogenous origin. Phosphatidylglycerol was found in all organs and tissues of the holothurians C. frondosa japonica and E. fraudatrix, as well as in the gastrointestinal tract of ascidians; its origin is unclear.     

Expression of amino acid transporter genes in developmental stages and adult tissues of Antarctic echinoderms

Epithelial cells in the body wall of adult and developmental stages of marine invertebrates absorb dissolved organic material directly from seawater. Despite over a century of study, little is known about the molecular biological mechanisms responsible for this transport process. Previous studies on embryonic and larval Antarctic echinoderms show that amino acid uptake could provide an important supplement of metabolic substrates. In the present study, partial cDNA sequences of 11 putative amino acid transporter genes were isolated from six species of Antarctic echinoderms including the Antarctic sea stars Acodontaster hodgsoni, Diplasterias brucei, Odontaster meridionalis, Odontaster validus, and Perknaster fuscus, and the Antarctic sea urchin Sterechinus neumayeri. Conserved domains of cDNA-deduced amino acid sequences characterized these genes as being members of a family of amino acid transporters (solute carrier family 6). Expression of these genes was detected throughout embryonic and larval development of two species that have contrasting developmental modes (A. hodgsoni: lecithotrophic; O. meridionalis: planktotrophic). In all six species studied, the expression of amino acid transporter genes was detected in tube feet and digestive organs of adult animals, demonstrating that members of a single amino acid transporter gene family are expressed during the entire life history of a marine invertebrate. The identification of these genes is an important step toward developing a mechanistic understanding of amino acid transport capacities in Antarctic marine invertebrates.     

Persistence of antibiotic-resistant and -sensitive Proteus mirabilis strains in the digestive tract of the housefly (Musca domestica) and green bottle flies (Calliphoridae)

Synanthropic flies have been implicated in the rapid dissemination of antibiotic-resistant bacteria and resistance determinants in the biosphere. These flies stably harbor a considerable number of bacteria that exhibit resistance to various antibiotics, but the mechanisms underlying this phenomenon remain unclear. In this study, we investigated the persistence of antibiotic-resistant bacteria in the digestive tract of houseflies and green bottle flies, using Proteus mirabilis as a model microorganism. One resistant strain carried the blaTEM and aphA1 genes, and another carried a plasmid containing qnrD gene. Quantitative PCR and 454 pyrosequencing were used to monitor the relative abundance of the Proteus strains, as well as potential changes in the overall structure of the whole bacterial community incurred by the artificial induction of Proteus cultures. Both antibiotic-resistant and -sensitive P. mirabilis strains persisted in the fly digestive tract for at least 3 days, and there was no significant difference in the relative abundance of resistant and sensitive strains despite the lower growth rate of resistant strains when cultured in vitro. Therefore, conditions in the fly digestive tract may allow resistant strains to survive the competition with sensitive strains in the absence of antibiotic selective pressure. The composition of the fly-associated bacterial community changed over time, but the contribution of the artificially introduced P. mirabilis strains to these changes was not clear. In order to explain these changes, it will be necessary to obtain more information about bacterial interspecies antagonism in the fly digestive tract.     

EVOLUTIONARY LOSS OF LARVAL FEEDING: DEVELOPMENT,FORM AND FUNCTION IN A FACULTATIVELY FEEDING LARVA,BRISASTER LATIFRONS

Species with large eggs and nonfeeding larvae have evolved many times from ancestors with smaller eggs and feeding larvae in numerous groups of aquatic invertebrates and amphibians. This change in reproductive allocation and larval form is often accompanied by dramatic changes in development. Little is known of this transformation because the intermediate form (a facultatively feeding larva) is rare. Knowledge of facultatively feeding larvae may help explain the conditions under which nonfeeding larvae evolve. Two hypotheses concerning the evolutionary loss of larval feeding are as follows: (1) large eggs evolve before modifications in larval development, and (2) the intermediate form (facultatively feeding larva) is evolutionarily short-lived. I show that larvae of a heart urchin, Brisaster latifrons, are capable of feeding but do not require food to complete larval development. Food for larvae appears to have little effect on larval growth and development. The development, form, and suspension feeding mechanism of these larvae are similar to those of obligate-feeding larvae of other echinoids. Feeding rates of Brisaster larvae are similar to cooccurring, obligate-feeding echinoid larvae but are low relative to the large size of Brisaster larvae. The comparison shows that in Brisaster large egg size, independence from larval food, and relatively low feeding rate have evolved before the heterochronies and modified developmental mechanisms common in nonfeeding echinoid larvae. If it is general, the result suggests that hypotheses concerning the origin of nonfeeding larval development should be based on ecological factors that affect natural selection for large eggs, rather than on the evolution of heterochronies and developmental novelties in particular clades. I also discuss alternative hypotheses concerning the evolutionary persistence of facultative larval feeding as a reproductive strategy. These hypotheses could be tested against a phylogenetic hypothesis.     

Cytological and enzyme-histochemical investigations on the digestive organs of Nautilus pompilius (Cephalopoda, Tetrabranchiata)

The foregut, stomach, caecum, midgut, and rectum of the digestive tract of Nautilus pompilius L.were investigated with ultrastructural and enzyme-cytological methods. Three different cell types were identified within the lamina epithelialis mucosae: main cells, goblet cells, and cells with secretory granules. The main cell type is the epithelial cell with microvilli, a basal nucleus surrounded by dictyosomes, rough endoplasmic reticulum, mitochondria, and electron-dense granules identified as lysosomes in the apical part of the cell. In the caecum this cell type contains endosymbiotic bacteria. The presence of endocytotic vesicles and the storage of lipids in the caecum indicate that this organ is involved in the process of absorption. In the caecum and the longitudinal groove of the rectum the main cells are, in addition, ciliated, facilitating the transport of food particles and faeces. Two types of goblet cells are found in all organs except in the stomach, forming a gliding path for food particles and protecting the epithelium. In the foregut and rectum, cells with electron-dense granules were recognized as the third type. The conspicuous secretory cells of the rectum represent a delimited rectal gland; its possible biological function is discussed. The tunica muscularis in all organs of the digestive tract consists of obliquely striated muscle cells innervated by axons containing transparent, osmiophilic and dense-cored vesicles. Positive reactions for acid and alkaline phosphatase, monoamine oxidase, β-glucuronidase, and trypsin- and chymotrypsin-like enzymes are localized in the lamina epithelialis mucosae.     

Morphological similarity between echinoid taxa as a result of convergent and parallel evolution

Convergence is observed in groups, which are phylogenetically remote. A flat test is typical for some representatives of the order Cassiduloida, such as Jurassic and Early Cretaceous species of the genera Pygurus and Clypeus, and also many Cenozoic «sand dollars,» i.e., echinoids of the order Clypeasteroida. Both usually inhabit coarse sandy grounds of shallow areas. The superorder Spatangacea includes the so-called Echinocorys life form, which is characterized by an oval test with superficial nonpetaloid or subpetaloid ambulacra, marginal or inframarginal periproct, and absence of fascioles. These are the following genera: Early Cretaceous Corthya (family Collyritidae), Late CretaceousLate Paleocene Echinocorys (family Holas-teridae), Paleocene Isaster, Recent Isopatagus (family Isasteridae), Recent Scrippsechinus (family Palaeotro-pidae), and Recent Urechinus (family Urechinidae). In contrast to the majority of spatangaceans with the burrowing mode of life, these genera dwell on the substrate surface. In the Cenozoic, the monobasal apical system appears in some genera of the order Spatangoida, most genera of the order Cassiduloida, all groups of the orders Clypeasteroida and Oligopygoida, and in the genus Echinoneus (order Holectypoida). The paral-lelism is revealed in groups connected by remote relationships. At the end of the Middle Jurassic (Callovian) and, especially, Late Jurassic, the so-called disasterid echinoids (superorder Spatangacea) show a distinct trend to the loss of contact between ocular plates I and V and apices of the posterior ambulacra with the periproct, which are shifted to the anterior part of the apical system (genera Collyrites, Collyropsis, Cyclolam-pas). At the same time, the peristome of some genera was displaced to the anterior margin of the test, which became bilaterally symmetrical in outline. However, in the Jurassic, all spatangaceans remained disasterid echinoids, i.e., had a disjunct apical system, which can be interpreted as a somewhat “abnormal” state. This trend disappeared only at the beginning of the Cretaceous, when “normal” forms with a joint apical system appeared, that is, the families Holasteridae (genera Eoholaster and Holaster, order Holasteroida) and Toxas-teridae (genus Toxaster, order Spatangoida). Interesting examples of synchronous parallelism are provided by the appearance of meridosternous (diasternal) plastron in two collyritid genera (Tetraromania and Corthya) in the Barremian, whereas in the holasterid genus Holaster, this type of plastron apparently appeared in the Valanginian (heterochronous parallelism). The ethmolitic type of the apical system appeared at the end of the Cretaceous and Paleocene at least in five families: Schizasteridae, Paleopneustidae, Brissidae, Spatangidae, and Loveniidae.     

中华稻蝗消化道内分泌细胞的鉴别与定位

采用整块组织Grimelius银染法和过氧化物酶标记的链霉亲和素免疫组织化学技术,结合生物统计学分析,对中华稻蝗Oxya chinensis消化道内分泌细胞进行鉴别与定位。结果表明：嗜银细胞分布于中华稻蝗的胃盲囊、中肠和后肠各段,以中肠和直肠中最多（P<0.05）, 前肠中未见分布。免疫组织化学法检测出了五羟色胺（5-hydroxytryptamine, 5-HT）、 胃泌素（gastrin, Gas）、 胰高血糖素（glucagon, Glu）和胰多肽（pancreatic polypeptide, PP）细胞, 未检出生长抑素（somatostatin, SS）细胞。免疫阳性细胞分布于中肠和后肠中, 前肠中未见分布。5-HT细胞和Gas细胞均主要分布于胃盲囊、中肠及直肠中,且均以直肠中最多（P<0.05）。Glu细胞在胃盲囊及整个中、后肠均有分布, 在中肠和直肠中最多（P<0.05）。PP细胞主要分布于中肠、回肠和直肠中,中肠中分布密度最大（P<0.05）。本研究显示中华稻蝗消化道中存在多种内分泌细胞,它们的分布情况与其他节肢动物相比存在一定的共性,也有其一定的特异性,可能与中华稻蝗特定的消化道结构和消化生理功能有关。     

Novel tetraamines,pentaamines and hexaamines in sea urchin,sea cucumber,sea squirt and bivalves

• 1.1. Polyamines were extracted from the guts and ovaries of the sea urchin Anthocidoris crassispina, and the guts and flesh of the sea cucumber Stichopus japonicus and the sea squirt Halocynthia roretzi, the oyster Crassostrea gigas and the short-necked clam Tapes philippinarum, and analyzed by ion-exchange high-performance liquid chromatography and gas chromatography-mass spectrometry.
• 2.2. Norspermidine and norspermine as well as putrescine, cadaverine, spermidine, spermine and agmatine were the ubiquitous polyamines in these invertebrates. These results suggest the widespread distribution of norspermidine and norspermine in invertebrates.
• 3.3. Thermopentamine, thermohexamine and homothermohexamine were found in the sea urchin. This in the first report on the occurence of thermopentamine and hexaamine in invertebrates.
• 4.4. Homospermidine, canavalmine, aminopropylhomospermidine, homospermine, caldopentamine, homocaldopentamine and aminopropylcanavalmine were found in the sea cucumber. Homospermidine, aminopropylhomospermidine and homospermine were found in the squirt. This is the first report on the occurence of canavalmine, aminopropylhomospermidine, homospermine, homocaldopentamine and aminopropylcanavalmine in invertebrates.

     

The clam Meretrix lamarckii (Bivalvia: Veneridae) is a rich repository of marine lactic acid bacterial strains

Lactic acid bacteria (LAB) were isolated from the intestinal tract of the wild clam Meretrix lamarckii caught from the coastal waters of Kashima, Ibaraki, Japan. As many as 415 isolates were obtained using the culture method, of which 70 were considered presumptive LAB strains based on phenotypic tests. Phylogenetic analysis of these presumptive isolates of LAB based on the sequence of the 16S rRNA gene demonstrated that the species belonged to several genera of Lactobacillus, Lactococcus and Pediococcus. Interestingly, however, the species composition was different between the samples in July and October 2010. Further analyses based on the fermentation profiles revealed that the LAB from the clam caught in July 2010 were identified to be Lactobacillus curvatus, Lactobacillus plantarum, Lactococcus lactis subsp. cremoris and Pediococcus pentosaceus, whereas those in October 2010 were identified to be Lactobacillus plantarum, Lactococcus lactis subsp. lactis and P. pentosaceus. The diversity of LAB in the intestinal tract of the clam suggests that the filter feeder bivalves such as M. lamarckii are a rich repository of marine isolates of LAB.     

Responses of the digestive tract of the omnivorous northern brown bandicoot, Isoodon macrourus (Marsupialia: Peramelidae), to plant- and insect-containing diets

The gastrointestinal tract of omnivores such as bandicoots (Marsupialia: Peramelidae) must be able to process foods as different as invertebrates, fungi and plant material. We studied the mechanisms involved in the utilisation by captive northern brown bandicoots (Isoodon macrourus) of insect larvae and milled lucerne (Medicago sativa) hay incorporated into a basal diet of a commercial small carnivore mix. Animals on the plant-basal mix digested less dry matter, energy, lipid, fibre and total nitrogen, but consumed 79% more dry matter than those on the insect-basal mix. Consequently intake of digestible energy (i.e. energy absorbed) was not significantly different between diets. Mean retention time (MRT, the mean time a marker remains in the tract) of a large particle marker was shorter on the plant-basal mix, reflecting its higher intake, but MRT of a solute marker was not significantly different between diets. Consequently the solute marker was retained longer than the particle marker on the plant-basal mix, indicating selective retention of solutes and very small particles in the caecum on this diet. This was confirmed by a higher proportion of small particles in the caecum than the distal colon of road-killed I. macrourus. Thus the main responses by I. macrourus to the plant-basal mix appeared to be an increase in gastrointestinal tract capacity (from radiographic evidence), selective retention of solutes and very small particles in the caecum, and facilitated passage of less tractable large particles through the colon. As a consequence, food intake was higher on the plant-basal mix, which compensated for its lower digestibility, and intake of digestible energy was similar to that on the insect-basal mix. This considerable flexibility of the morphologically rather simple digestive tract of northern brown bandicoots helps to explain their ability to cope with naturally variable diets consisting of mainly invertebrates in summer to much more plant and fungal material in winter, and to survive in nutritionally dynamic environments such as heathlands where there can be dramatic changes in food type and availability following periodic wildfires. Accepted: 14 April 1999