Spine Types Research Articles

Slices Have More Synapses than Perfusion-Fixed Hippocampus from both Young and Mature Rats

Hippocampal slices have long been used to investigate properties of synaptic transmission and plasticity. Here, for the first time, synapses in slices have been compared quantitatively with synapses occurring in perfusion-fixed hippocampus, which is presumed to represent the natural in vivo state. Relative to perfusion-fixed hippocampus, a remarkable 40-50% increase in spine number occurs in adult hippocampal slices, and a 90% increase occurs in slices from postnatal day 21 rats. Serial EM shows that all of the dendritic spines have normal synapses with presynaptic and postsynaptic elements; however, not all spine types are affected uniformly. Stubby and mushroom spines increase in the adult slices, and thin, mushroom, and branched spines increase in the immature slices. More axonal boutons with multiple synapses occur in the slices, suggesting that the new synapses form on preexisting axonal boutons. The increase in spine and synapse number is evident within a couple of hours after preparing the slices. Once the initial spine induction has occurred, no further change occurs for up to 13 hr in vitro, the longest time investigated. Thus, the spine increase is occurring during a period when there is little or no synaptic activity during the first hour, and the subsequent stabilization in spine synapse numbers is occurring after synaptic activity returns in the slice. These findings suggest that spines form in response to the loss of synaptic activity when slices are removed from the rest of the brain and during the subsequent 1 hr recovery period.

A new inaperturate pollen genus from the Late Miocene sediments of Site 218 of DSDP Leg 22 in the Central Bengal Fan, Indian Ocean

A new pollen genus Varisculptinaperturites has been described from the Late Miocene sediments of Site 218 of DSDP Leg 22 in the Central Bengal Fan, Indian Ocean. The genus accommodates inaperturate pollen ornamented with two types of sculpture, smaller spines and gemmae and larger clavae/baculae.

Mouthpart sensilla of cave species of australian paratemnopteryx cockroaches (BLATTARIA : BLATTELLIDAE)

The external morphology and distribution of sensilla on maxillary and labial palpi are described for 3 Australian cave-dwelling cockroach species, Paratemnopteryx stonei, P. howarthi, and P. sp. nov (Blattaria : Blattellidae), using scanning electron microscopy. The apex of the maxillary palpus expands ventrally into a flexible pad with a distinct sensillar field. The total number of sensilla ranges from 1045 (males)–1486 (females). Sensillar fields of females are larger and contain more sensilla than those of the males, although the density of sensilla is usually equal or greater for males. Papilla-like groove-and-slit chemoreceptors comprise 93–97% of the sensilla. Two types of trichoid sensilla occur along with bifurcuted sensilla and rare basiconic sensilla that are poreless and have an inflexible socket. Chaetic sensilla occur on all species and several thousand pectinate scales, with bulbous bases, and spicules are present in large fields on each side of the sensillar field. The apex of the labial palpus is only slightly expanded into a sensillar field. Sensillar diversity is similar to the maxillary palpus but there are only 5–29% as many sensilla, the number ranging from 97 (males)–346 (females). Sensillar fields of females are slightly to much larger than those of males but the sensillar density is equal between the sexes or greater on males. Two types of trichoid and one or two types of basiconic sensilla occur on each species. Chaetic chemoreceptors are present along with three types of non-socket spines. Sensilla varied moderately in morphology and gender-related abundance from those on epigean cockroach species.

Various types of suprameatal spines and depressions in the human temporal bone.

Since it covers the lateral wall of the mastoid air system, the suprameatal triangle is of importance to otologic surgeons during mastoidectomy. Because of this clinical importance, topographic anatomy of the suprameatal spine and depression was studied on Anatolian skulls. In all, 363 male and 231 female skulls were studied. The most prevalent type of suprameatal spine resembled a crest and was found in both sexes on the right (77.6%) and left (80%) sides. The absence of a suprameatal depression was significantly higher in females (right 9.1%; left 8.7%) than in males (right 1.7%; left 2.5%). Suprameatal depressions were mostly shallow in female subjects, but were mostly observed in males to be at a medium depth or deep.

Description and comparison of major foregut ossicles in hydrothermal vent crabs

Major foregut (gastric mill) ossicles, including the dorsal median tooth, lateral teeth, accessory lateral teeth, and cardiopyloric valve, of hydrothermal vent crabs were dissected and examined during the summer of 1996 from specimens housed at the Natural History Museum of Los Angeles County. Ossicles are described for two species of hydrothermal vent crabs (family Bythograeidae Williams, 1980). The western Pacific Austinograea williamsi Hessler and Martin has an unusual dorsal median tooth. The surrounding cuticular flange is scalloped and bears spinulose setae at the tip of each of the protruding edges, a condition perhaps unique in the Brachyura. The lateral teeth are mostly unremarkable, bearing the typical large anterior denticles and deep serrations seen in other crab families, but with a higher number of serrations than is known for any species previously described. The accessory lateral teeth bear flattened, plate-like spines that are widest basally and that taper to a cylindrical tip. The basic armature of the foregut of Bythograea thermydron Williams, known only from vents in the eastern Pacific, is very similar. Scalloping of the median tooth borders is less pronounced, however, and the shape of the tooth itself and of the plate from which it arises is slightly different. The lateral teeth bear fewer and more widely spaced grooves, and the cardiopyloric valve entrance appears less setose at its extremity. Comparison with foregut ossicles in other crab families based on earlier studies, most of which have not employed SEM, reveals some similarities between bythograeids and some xanthids, but does not clarify the phylogenetic position of the bythograeids. Because of the paucity of other SEM studies of the brachyuran foregut, it is difficult to ascertain whether some of the many spine and setal types in the bythograeid foregut are unique or even unusual compared to those of other crab families. Nothing about the foregut of the vent crabs is indicative of their unusual habitat. Anecdotal observations of feeding in vent crabs indicate that they are opportunistic scavengers and omnivores, which is in keeping with the non-specialized nature of the foregut. The debate between adaptation vs phylogeny as determinants of the form of the gastric mill components is briefly discussed.

THREE SPECIES OF THE DIATOM GENUS CORETHRON CASTRACANE: STRUCTURE, DISTRIBUTION AND TAXONOMY

Three species of the marine planktonic diatom genus Corethron are recognized, and compared in detail using light and electron microscopy. The confusion in the use of some of the names given to taxa in the past is clarified, mainly based on differences in the morphology of the hooked spines. The commonly used combination, Corethron criophilum Castracane is discarded in favour of the earlier described entity, Actiniscus pennatus which was combined with Corethron in 1903 as Corethron pennatum (Grunow) Ostenfeld. The function of the tfiree types of spine found in the genus is discussed. It is proposed that the spines of the two species existing as single cells, C. pennatum and C. hystrix, serve to move apart and maintain separation of the cells in the ocean. Similar spines are found in C. inerme but there the cells are normally joined by linking spines to form long chains. The hooked spines have a similar role to play but we suggest that their main function is to secure the developing spine system in order to safeguard the cingulum until the new valves reach maturity.

GENETIC VARIATION AND POLYMORPHISM IN THE INDUCIBLE SPINES OF A MARINE BRYOZOAN.

Of particular value in understanding the evolution of genotypes with broad phenotypic ranges (phenotypic plasticity) are the few examples of organisms with adaptive plasticity, such as those that develop a defensive morphology in response to cues from predators. We know little about the heritability of inducible defensive characters or the range of phenotypes available for selection to act on in the field. Membranipora membranacea is a colonial marine bryozoan that produces spines within two days of exposure to waterborne predator extracts. Surveys done in 1993 and 1995 showed that the population at Friday Harbor Labs, Washington, was polymorphic for inducible spine type and was composed of a constitutively spined type that produced spines in the absence of a predator stimulus, an unspined phenotype that did not produce spines irrespective of a stimulus, and an inducibly spined phenotype that would produce spines if exposed to the appropriate cue. In 1995, the frequencies of these types were determined through a laboratory common-garden experiment; 178 colonies were cultured from metamorphosis through approximately 30 days and then exposed to the cue. The inducible type was the dominant, comprising 80.3% of the population. The constitutive type made up 6.2% of the population, and the remaining 13.4 % was the unspined type. The frequency of the three types was similar to a preliminary trial of the experiment run in 1993. Experiments also showed that the lengths of the spines of the inducible type varied continuously among genotypes. To assess causes of variation in the inducible spine response and its clonal heritability, 16 clones were subdivided and grown in a common environment and exposed to a single dosage of spine inducing substance (SIS). Spine length showed high clonal heritability. The range of colony responses from a single environment varied from relatively unresponsive to highly responsive colonies with a very low threshold of response. Norms of reaction were quantified for spine lengths of inducible genotypes originating from two field environments by testing them in a concentration series of SIS. Both spine length and spine type varied with concentration of inducer. Within a clone, colonies were more likely to produce membranous spines than corner spines at higher concentrations. At low concentrations, only straight spines were produced. This study showed that populations of M. membranacea at Friday Harbor are a mix of inducible, nonspined and constitutively spined individuals. Even the inducible individuals showed high heritable variation in the length of spine activated, suggesting that there is considerable scope for the evolution of this character. A norm-of-reaction experiment further showed that the type of spine produced, membranous or corner, varied with the concentration of the cue. Factors maintaining the polymorphism and the broad range of genotypes could include high costs of defending the spined types coupled with a shifting biotic regime.

<i>Ulex parviflorus sensu lato (Genisteae, Leguminosae)</i> en la zona centro de Portugal

A group of gorses growing in Central Portugal has been traditionally identified as Ulex parviflorus Pourret sensu lato . The study of these plants reveals important differences in morphological characters such as size, ramification, colour and indumentum of shoots, the type, disposition and indumentum of primary and secondary spines, and the dimensions, shape and indumentum of the calyx. Based on these morphological characteristics, as well as chromosome number, ecological requirements and geographic distribution, the plants are now ascribed to the following taxa: U. airensis sp. nov., U. australis subsp. welwitschianus stat. & comb. nov., U. eriocladus C. Vicioso and U. jussiaei Webb

Australodiscus peruvianus, gen. et sp. nov., a marine centric diatom from the Peruvian Eocene

A new diatom genus and species, Australodiscus peruvianus Porguen et Sullivan, is described from an Eocene marine deposit from the Paracas Peninsula, Peru. The valve outline is stellate with ocelli and rimoportulae situated singly on alternate points of the star. The rimoportulae are winged and situated on a secondary marginal ridge. Three types of spines are present; two of these are also winged. Our new form appears to be best placed for the present in the centric family Triceratiaceae (Schütt) Lemmermann, being most closely related to the type genus Triceratium Ehrenberg and Grovea A. Schmidt.

Australodiscus peruvianus, gen. et sp. nov., a marine centric diatom from the Peruvian Eocene

A new diatom genus and species, Australodiscus peruvianus Porguen et Sullivan, is described from an Eocene marine deposit from the Paracas Peninsula, Peru. The valve outline is stellate with ocelli and rimoportulae situated singly on alternate points of the star. The rimoportulae are winged and situated on a secondary marginal ridge. Three types of spines are present; two of these are also winged. Our new form appears to be best placed for the present in the centric family Triceratiaceae (Schütt) Lemmermann, being most closely related to the type genus Triceratium Ehrenberg and Grovea A. Schmidt.

Integrated facility layout and material handling system design in semiconductor fabrication facilities

Semiconductor manufacturing is an important component of the U.S. manufacturing industry. Most of today's fabrication facilities and those being designed for the near future use a bay layout configuration and an overhead monorail system for moving material between bays. These material handling systems are usually designed with a spine or perimeter type of configuration. This paper investigates the layout and material handling system design integration problem in semiconductor fabrication facilities and proposes a methodology for solving this integrated design problem. A spacefilling curve approach is used to address the facility layout, while the structure of the spine and perimeter configurations are exploited to create a network flow problem to determine the material handling system design. Computational results are presented and show exceptional promise for this procedure in solving the integrated design problem in a semiconductor manufacturing environment.

Estradiol increases the frequency of multiple synapse boutons in the hippocampal CA1 region of the adult female rat.

The effect of estradiol to increase the density of dendritic spines and axospinous synapses on hippocampal CA1 pyramidal cells in the adult female rat has been well-documented. However, presynaptic involvement in this process of synapse elimination and formation in the adult is unknown. To address this issue, we have reconstructed 410 complete presynaptic boutons through coded serial electron micrographs of CA1 stratum radiatum to determine the: (1) frequency of multiple (MSB) vs. single (SSB) synapse boutons; (2) number of synaptic contacts per MSB; (3) bouton volume and surface area; and (4) types of spines in synaptic contact with MSBs and SSBs in ovariectomized, estradiol-treated animals (OVX + E) versus ovariectomized oil-treated controls (OVX + O). Quantitative analysis of this tissue revealed that, in OVX + E animals, 45.0% of presynaptic boutons form multiple synaptic contacts with dendritic spines compared to 27.3% in controls (P < 0.01); the average number of synapses per dendritic spines compared to 27.3% in controls (P < 0.01); the average number of synapses per MSB was 2.7 in OVX + E animals compared to 2.3 in controls (P < 0.05). This represents a 25.5% increase in the number of synapses formed by a given number of presynaptic boutons in estradiol-treated animals (P < 0.01) which largely accounts for the previously observed estradiol-induced increase in axospinous synapse density. There was no treatment effect on bouton size; however, because MSBs are larger than SSBs, the increased frequency of MSBs in estradiol-treated tissue results in a trend toward an estradiol-induced increase in average bouton size. Additionally, MSBS were found to be more irregular in shape, i.e., significantly less spherical, than SSBs. Our results indicate that estradiol-induced dendritic spines form synapses primarily with preexisting boutons in stratum radiatum and that these boutons enlarge and change shape as they accommodate new synapses. Such findings suggest a relatively active role for dendrites in the process of adult synapse formation.

Estradiol increases the frequency of multiple synapse boutons in the hippocampal CA1 region of the adult female rat

The effect of estradiol to increase the density of dendritic spines and axospinous synapses on hippocampal CA1 pyramidal cells in the adult female rat has been well-documented. However, presynaptic involvement in this process of synapse elimination and formation in the adult is unknown. To address this issue, we have reconstructed 410 complete presynaptic boutons through coded serial electron micrographs of CA1 stratum radiatum to determine the: (1) frequency of multiple (MSB) vs. single (SSB) synapse boutons; (2) number of synaptic contacts per MSB; (3) bouton volume and surface area; and (4) types of spines in synaptic contact with MSBs and SSBs in ovariectomized, estradiol-treated animals (OVX + E) versus ovariectomized oil-treated controls (OVX + O). Quantitative analysis of this tissue revealed that, in OVX + E animals, 45.0% of presynaptic boutons form multiple synaptic contacts with dendritic spines compared to 27.3% in controls (P < 0.01); the average number of synapses per MSB was 2.7 in OVX + E animals compared to 2.3 in controls (P < 0.05). This represents a 25.5% increase in the number of synapses formed by a given number of presynaptic boutons in estradiol-treated animals (P < 0.01) which largely accounts for the previously observed estradiol-induced increase in axospinous synapse density. There was no treatment effect on bouton size; however, because MSBs are larger than SSBs, the increased frequency of MSBs in estradiol-treated tissue results in a trend toward an estradiol-induced increase in average bouton size. Additionally, MSBs were found to be more irregular in shape, i.e., significantly less spherical, than SSBs. Our results indicate that estradiol-induced dendritic spines form synapses primarily with preexisting boutons in stratum radiatum and that these boutons enlarge and change shape as they accommodate new synapses. Such findings suggest a relatively active role for dendrites in the process of adult synapse formation. © 1996 Wiley-Liss, Inc.

Variability and Grouping of Northwestern Spanish Chestnut Cultivars. I. Morphological Traits

Different chestnut (Castanea sativa Mill.) cultivars are at present grown in the region of Galicia, northwestern Spain, but no distinguishing traits among cultivars have been defined so far. The objectives of this research were to 1) describe the intra and intercultivar variability of chestnut cultivars; 2) define primary morphological traits to be useful for a simple morphological classification system of the cultivars; and 3) study the association between some environmental variables and the morphological traits. Seventeen morphological traits in a sample of 373 trees belonging to 82 local cultivars of chestnut were studied by methods of numerical taxonomy, principal component and cluster analysis. These traits were selected from 135 previously studied as having possible discriminating taxonomic value. Significant variability among cultivars and among trees within cultivars was found for most of the traits. The trees were grouped according to the degree of dissimilarity on the basis of the Mahalanobis generalized distance. Most of the clones collected under a specific cultivar name were included within the same cluster group enabling us to classify 53 of the cultivars studied. A hierarchical classification system that identifies eight cultivar groups is proposed based on four discriminating levels: nut size, fruit shape, male flower type and length of burr spines. Most of the correlations between the environmental variables and the morphological traits were no significant or had a low value. The lack of correlation between the environmental variables and nut size indicates that this important trait is under strong genetic control, it is not influenced by environmental conditions and it is consistent throughout the area sampled.

Structurally variant classes of pilus appendage fibers coexpressed from Burkholderia (Pseudomonas) cepacia

One or more of five morphologically distinct classes of appendage pili were determined to be peritrichously expressed by Burkholderia (formerly Pseudomonas) cepacia isolated from disparate sources. B. cepacia-encoded cblA pilin gene hybridization-based analysis revealed that one associated class, cable (Cbl) adhesin type IIB. cepacia pili, correlates with epidemically transmitted strains from a single cystic fibrosis (CF) center. When only phenotypic assays were available, correlations between the source and the pilus type were nonetheless observed: filamentous (Fil) type IIIB. cepacia pili correlated with CF-associated nonepidemic isolates, spine (Spn) type IVB. cepacia pili correlated with clinical (non-CF) isolates, and spike (Spk) type VB. cepacia pili correlated with environmental isolates. Further, Cbl, Fil, or Spk pili typically appear as an internal framework for constitutively coexpressed, peritrichously arranged dense mats of fine, curly mesh (Msh) type IB. cepacia pili. Constitutive coexpression of dense mats of Msh type IB. cepacia pili in association with a labyrinth of either Cbl, Fil, or Spk pili suggests possible cooperative pilus interactions mediating adhesion-based colonization in the differing environments from which the strains were isolated. Despite such correlations, phylogenetic analyses indicate that with the exception of the epidemically transmitted clusters of isolates, the remaining B. cepacia strains from the other three sources exhibited an equal degree of genetic relatedness independent of origin. As previously found for Escherichia coli, this discrepancy could be accounted for by selection-driven, in vivo horizontal transfer events between distantly related members of the species B. cepacia, leading to the genetic acquisition of environmentally appropriate adhesion-based colonization pilus operons.

Spine morphology of neurons in the avian forebrain is affected by rearing conditions

An area of the caudal forebrain of male zebra finches, the Archi-Neostriatum caudale (ANC), which is active during arousal (Bischof & Herrmann, 1986, 1988), shows rearing-dependent changes in neuron morphology (Rollenhagen & Bischof, 1991). We demonstrate here that rearing conditions also affect the shape of spines of one of the four ANC neuron types. This neuron type was examined in birds reared under five different conditions--in isolation (1), caged (2), in the aviary (3), and with social contact (4) or chasing (5) after an isolation period. Our results show that social experience determines the proportion of the three types of spines (thin, mushroom, and stubby) of the investigated neuron type. Rearing conditions and short social contact also affect the spine stem length of the thin spine type. Long-term isolation results in a reduction in number and elongation of shafts of thin spines, along with an increase of stubby-and mushroom-shaped spines. Short-term social contact or arousal enhances the number of mushroom-and thin-shaped spines and reduces the length of spine stems of thin spines. We suggest that isolation prevents the ANC neuron from reaching full development. The increase of mushroom and thin spine types due to social contact indicates that the stubby-shaped spines are replaced by, or transformed into, mushroom-shaped spines, and the mushroom-shaped spines are replaced by, or transformed into, thin spines. These results confirm and extend the experimental background for our hypothesis (Rollenhagen & Bischof, 1991) that social contact is necessary for development of normal morphology of ANC neurons.

Scanning electron microscopy of the cuticular armature of the nematode Gnathostoma spinigerum Owen, 1836 from cats in Laos

The body surface of an immature female Gnathostoma spinigerum found for the first time in the definitive host (Felis catus f. domestica) in Laos was studied using a scanning electron microscope. All types of cuticular spines, which are one of the most important features for species identification of gnathostomid nematodes, together with their spatial arrangement, are described and figured.

Classification of Screwworms (Diptera: Calliphoridae) by Larval Spine Morphology

The morphology of screwworm, Cochliomyia hominivorax (Coquerel), larval spines was affected by larval substrate and geographic effects on variation. Spine morphology of individual larvae was used to determine criteria for discrimination analysis classification. Classification criteria for types of wounds and host species were determined for larvae from central and southern Mexico. Larvae from marginal populations (United States, Caribbean Islands, Costa Rica) were classified by these criteria. Classifications were 27% accurate for wound type and 20% correct for animal species. When single egg masses were split and larvae were reared on laboratory diet and animal substrates, analysis of spine types showed significant rearing environment and sibship (egg mass) effects. Cluster analysis of geographic groups showed that larvae from central populations were closely clustered, whereas marginal groups were morphologically dissimilar from each other and from the central populations. These results indicate that larval spine morphology has limited value in taxonomic or systematic studies of screwworms and that mating compatibility was not related to spine similarity among populations. The significant differences between wound-reared and artificial media-reared larvae indicate that differences in spine morphology could be used to verify that samples from the field were indeed taken from animal wounds.

Ultrastructural organization of normal and transplanted rat fascia dentata: I. A qualitative analysis of intracerebral and intraocular grafts.

Few studies have dealt with the general ultrastructure and synaptic organization of grafted brain tissue. This study was therefore performed to extend current light microscopic observations on intracerebral and intraocular grafts of hippocampal tissue to the ultrastructural level. Blocks of tissue containing the hippocampus and fascia dentata from day 21 embryonic rats were grafted into the brain of developing and adult rats and to the anterior eye chamber of adult rats. After 100 or 200 days of survival the recipient rat brains or eyes were processed for electron microscopy. Tissue containing the graft dentate molecular layer and adjacent granule cell layer was selected for ultrastructural analysis, together with a few samples of the hilus and CA3. Normal dentate tissue was analyzed as control. At the light microscopic level most intracerebral and intraocular grafts displayed an organotypic organization with clearly recognizable cell and neuropil layers. Under the electron microscope the grafted granule cells had normal-appearing dendrites bearing the normal types of spines and forming the normal types of synapses. This was the case even in the absence of the normal major extrinsic afferents like the perforant path. The graft dentate granule cells formed axons and terminals with characteristics of the normal mossy fiber system in the hilus and CA3, in addition to aberrant supragranular mossy fiber terminals known from light microscopic studies of dentate transplants. Abnormal structures included a few dendritic growth cones and an increased occurrence of polyribosomes in spines. Their occurrence indicates ongoing dendritic plasticity even 100 days after transplantation. There was also an increased density of glial elements, particularly in the intraocular grafts. In some of these grafts the granule cells displayed immature traits in terms of nuclear indentations. Dentate interneurons of the basket cell type were present in both the intracerebral and the intraocular grafts. We conclude that grafted dentate granule cells, in different surroundings and without the normal, major perforant path input, can develop a basically normal cellular morphology, which includes the normal ultrastructural characteristics of the dendrites with spines and synapses, and the mossy fibers and its terminals.

Frontal ambulacral and peribuccal areas of the spatangoid echinoid Echinocardium cordatum (Echinodermata): a functional entity in feeding mechanism

Echinocardium cordatum (Pennant, 1777) were collected in Normandy, France, in March and November 1985. The grooved frontal ambulacrum of the burrowing echinoid E. cordatum functions in transferring surface sediment from the apex to the mouth of the echinoid. Particles that fall down the burrow's chimney accumulate between the spines of the apical tuft and are taken over by specialized groove spines. Due to the slope of the groove and the type of floor spines it harbours, four successive regions can be recognized, namely the adapical region, the fasciolar region, the ambital region, and the adoral region. As a general rule aboral floor spines (i.e. club-shaped, golf-club-shaped, and isodiametric spines) function in gathering particles while propelling them mouthwards; they simultaneously embed particles in a mucous string. The oral floor spines (i.e. spatulated spines) function in hauling and guiding the mucous string towards the peribuccal area of the echinoid. Once facing the peribuccal area, particles are picked up by the peribuccal tube feet that either transfer them into the esophagus or scrape them out against the peribuccal spines. Spine-retained particles are either sucked up by the pumping action of the esophagus or fall to the burrow's floor where the tube feet may pick them up again. Together the apical tuft, the frontal groove, and the peribuccal area form an efficient food-collecting system that transfers trapped surface particles from the echinoid apex down to the mouth. Compared to that of most other spatangoids the frontal ambulacrum of E. cordatum is highly specialized. Such specialization is related to colonization of fine sediment, as is the occurrence of a chimney linking the burrow to the sediment surface. Actually the burrow's chimney is the only route for oxygen and food supply; it acts as a real ecological umbilical cord for spatangoids buried in fine sediment.