Modeling the role of incisures in vertebrate phototransduction

Phototransduction is mediated by a G-protein-coupled receptor-mediated cascade, activated by light and localized to rod outer segment (ROS) disk membranes, which, in turn, drives a diffusion process of the second messengers cGMP and Ca2+ in the ROS cytosol. This process is hindered by disks-which, however, bear physical cracks, known as incisures, believed to favor the longitudinal diffusion of cGMP and Ca2+. This article is aimed at highlighting the biophysical functional role and significance of incisures, and their effect on the local and global response of the photocurrent. Previous work on this topic regarded the ROS as well stirred in the radial variables, lumped the diffusion mechanism on the longitudinal axis of the ROS, and replaced the cytosolic diffusion coefficients by effective ones, accounting for incisures through their total patent area only. The fully spatially resolved model recently published by our group is a natural tool to take into account other significant details of incisures, including their geometry and distribution. Using mathematical theories of homogenization and concentrated capacity, it is shown here that the complex diffusion process undergone by the second messengers cGMP and Ca2+ in the ROS bearing incisures can be modeled by a family of two-dimensional diffusion processes on the ROS cross sections, glued together by other two-dimensional diffusion processes, accounting for diffusion in the ROS outer shell and in the bladelike regions comprised by the stack of incisures. Based on this mathematical model, a code has been written, capable of incorporating an arbitrary number of incisures and activation sites, with any given arbitrary distribution within the ROS. The code is aimed at being an operational tool to perform numerical experiments of phototransduction, in rods with incisures of different geometry and structure, under a wide spectrum of operating conditions. The simulation results show that incisures have a dual biophysical function. On the one hand, since incisures line up from disk to disk, they create vertical cytoplasmic channels crossing the disks, thus facilitating diffusion of second messengers; on the other hand, at least in those species bearing multiple incisures, they divide the disks into lobes like the petals of a flower, thus confining the diffusion of activated phosphodiesterase and localizing the photon response. Accordingly, not only the total area of incisures, but their geometrical shape and distribution as well, significantly influence the global photoresponse.     

Distinct claudins and associated PDZ proteins form different autotypic tight junctions in myelinating Schwann cells

The apposed membranes of myelinating Schwann cells are joined by several types of junctional specializations known as autotypic or reflexive junctions. These include tight, gap, and adherens junctions, all of which are found in regions of noncompact myelin: the paranodal loops, incisures of Schmidt-Lanterman, and mesaxons. The molecular components of autotypic tight junctions have not been established. Here we report that two homologues of Discs Lost-multi PDZ domain protein (MUPP)1, and Pals-associated tight junction protein (PATJ), are differentially localized in myelinating Schwann cells and associated with different claudins. PATJ is mainly found at the paranodal loops, where it colocalized with claudin-1. MUPP1 and claudin-5 colocalized in the incisures, and the COOH-terminal region of claudin-5 interacts with MUPP1 in a PSD-95/Disc Large/zona occludens (ZO)-1 (PDZ)-dependent manner. In developing nerves, claudin-5 and MUPP1 appear together in incisures during the first postnatal week, suggesting that they coassemble during myelination. Finally, we show that the incisures also contain four other PDZ proteins that are found in epithelial tight junctions, including three membrane-associated guanylate-kinase proteins (membrane-associated guanylate-kinase inverted-2, ZO-1, and ZO-2) and the adaptor protein Par-3. The presence of these different tight junction proteins in regions of noncompact myelin may be required to maintain the intricate cytoarchitecture of myelinating Schwann cells.     

Presence of the myelin-associated glycoprotein correlates with alterations in the periodicity of peripheral myelin

The myelin-associated glycoprotein (MAG) is an integral membrane protein (congruent to 100,000 mol wt) which is a minor component of purified peripheral nervus system (PNS) myelin. In the present study, MAG was localized immunocytochemically in 1-micrometer thick Epon sections of 7-d and adult rat peripheral nerves, and its localization was compared to that of the major structural protein (Po) of PNS myelin. To determine more precisely the localization of MAG, immunostained areas in 1 micrometer sections were traced on electron micrographs of identical areas from adjacently cut thin sections.l MAG was localized in periaxonal membranes. Schmidt-Lantermann incisures, paranodal membranes, and the outer mesaxon of PNS myelin sheaths. Compact regions of PNS myelin did not react with MAG antiserum. The results demonstrate MAG's presence in "'semi-compact" Schwann cell or myelin membranes that have a gap of 12-14 nm between extracellular leaflets and a spacing of 5 nm or more between cytoplasmic leaflets. In compact regions of the myelin sheath which do not contain MAG, the cytoplasmic leaflets are "fused" and form the major dense line, whereas the extracellular leaflets are separated by a 2.0 nm gap appearing as paired minor dense lines. Thus, it is proposed that MAG plays a role in maintaining the periaxonal space, Schmidt-Lantermann incisures, paranodal myelin loops, and outer mesaxon by preventing "complete" compaction of Schwann cell and myelin membranes. The presence of MAG in these locations also suggests that MAG may serve a function in regulating myelination in the PNS.     

Light microscopy study of Schmidt-Lantermann incisures in the guinea pig sciatic nerve

The authors studied the shape, orientation and number of Schmidt-Lantermann incisures in the sciatic nerve of the healthy guinea pig. The nerves were removed, fixed, shredded and evaluated in the light microscope. Internode length, fibre diameter and segment length were measured in isolated nerve fibres with an eyepiece micrometer. The number and configuration of the Schmidt-Lantermann incisures in the internode were found to be variable. The incisures follow one another at irregular intervals, dividing the whole internode into cylindroconical segments of varying lengths. The measured values were processed on a PMD 85/2 personal microcomputer.     

Some studies on Tylenchorhynchinae: proposal ofDivittus n.g. (Nematoda: Tylenchida)

Summary A new genusDivittus is proposed in the subfamily Tylenchorhynchinae Eliava, 1964 with only three incisures in the non-areolated lateral fields and the body cuticle without longitudinal ridges or lamellae. It is also proposed to restrictTrilineellus Lewis & Golden, 1981 to its type species only, i.e.,T. clathrocutis which has three incisures in the lateral fields and longitudinal ridges or lamellae on the body, andTylenchorhynchus to species which have four incisures in the lateral fields but no ridges or lammellae.T. divittatus Siddiqi, 1961,T. triglyphus Seinhorst, 1963 andT. sculptus Seinhorst, 1963 which were transferred toTrilineellus by Lewis & Golden (1981) are now included inDivittus with the following species ofTylenchorhynchus which have three incisures in the lateral fields:T. obscurisulcatus Andrássy, 1959,T. chonai Sethi & Swarup, 1968,T. madrasensis Gupta & Uma, 1981 andT. pruni Gupta & Uma, 1981. The synonymy ofT. chonai withT. triglyphus proposed by Tarjan (1973) is not accepted.Divittus labiatus n.sp. from soil around roots of sugarcane from north-eastern Nigeria is described. It is characterized by having the lip region continuous with the body, the spear 14–16 m long, the basal oesophageal bulb with a small overlap over the intestine and the tail with a large post-anal intestinal sac and 20 to 30 annules. A key to species ofDivittus is provided. ac]19830105     

Descriptions of Two New Species of Chronogaster Cobb, 1913 from India

Two new species of Chronogaster in India were described and illustrated, based on light and scanning electron microscopy. Chronogaster neotypica n. sp. collected from a sewage slurry was characterized by a medium-sized body, a ventral tail mucro without additional spines, absence of longitudinal incisures in lateral fields, and by the presence of crystalloids in the body. Diagnostic for C. spinicauda n. sp. collected from soil around roots of mango were a medium-sized body, a tail mucro with 10 spines, and absence of lateral lines and crystalloids. Males were not found.     

The taxonomy of Tylenchorhynchinae (Nematoda: Tylenchida) with longitudinal lines and ridges

Summary The taxonomy of those Tylenchorynchinae which have longitudinal lines or ridges on the cuticle is discussed. Dolichorhynchus is restricted to species with four incisures in the lateral field, lateral vulval flaps and a terminally notched bursa. D. prophasmis n. sp. is described. Neodolichorhynchus n. g. is erected for those species previously in Dolichorhynchus which have no lateral vulval flaps and a normal bursa, including: N. microphasmis (Loof, 1959) n.comb., N. judithae (Andrássy, 1962) n.comb., N. sulcatus (de Guiran, 1967) n.comb., and N. gladiolatus (Fortuner & Amougou, 1973) n.comb. Tessellus n.g. is proposed for T. claytoni (Steiner, 1937) n.comb. and T. pachys (Thorne & Malek, 1968) n.comb. the only remaining Tylenchorhynchus species with longitudinal cuticular lines. They are characterized by a rounded non-offset lip region, four incisures in the lateral field and a cuticular annulation divided into prominent blocks by deep longitudinal cuticular lines.     

Morphology of Schmidt-Lanterman incisures of human shoulder plexus nerves

Morphological studies of peripheral human nerves have shown that the number o of Schmidt-Lanterman incisures in an internode is proportional to the fibre diameter. Quantitative and ultrastructural examinations of incisures in the shoulder plexus nerves of the juvenile have proved some common features in the distribution and structure of incisures, characteristic for the nerves of both mammals and people. However, in all probability, in the peripheral nerves of man under certain condition there is a mechanism, leading to a sharp increase of the number of incisures in the internodal segment.     

Oriented spermatozoa in the spermatophore of the palaemonid shrimp, Macrobrachium rosenbergii

Estimates of the degree of orientation of spermatozoa within spermatophores of the freshwater shrimp, Macrobrachium rosenbergii, were made using electron micrographs of ultrathin sections cut transverse and parallel to the longitudinal axis of the spermatophores. Counts of the total number of longitudinal and non-longitudinal profiles of sperm spikes in transverse sections of spermatophores indicated that 67% of the spermatozoa were oriented with their spikes approximately perpendicular to the long axis of the spermatophore. In longitudinal sections, ~30% of the sperm spike profiles were oriented parallel to the longitudinal axis of the spermatophore, leaving ~70% of the profiles oriented in planes not parallel to the longitudinal spermatophore axis. Stereologic analyses based on the number of intersections of longitudinally sectioned sperm spike profiles per unit length of test lines placed over photographic images of ultrathin sections cut in both transverse and longitudinal planes of spermatophores gave a Saltykov approximation of 0.607, favoring the view that the spermatozoa were oriented with their spikes perpendicular to the long axis of the spermatophore. This orientation brings many of the sperm bases approximately parallel to the surface of the spermatophore, an orientation which may facilitate normal sperm-egg interaction during fertilization in this species.     

Ditylenchus destructor in Hulls and Seeds of Peanut

The potato rot nematode, Ditylenchus destructor Thorne, is reported for the first time in hulls and seeds of peanut. The populations found differed from D. dipsaei and D. myceliophagus in habitat, number of lateral incisures, shape of tail tip, and length of postvulval sac. Infected hulls had brown necrotic tissue at the point of connection with the peg, and a black discoloration appeared first along the longitudinal veins. Infected seeds were usually shrunken, and testae and embryos had a yellow to brown or black discoloration. Of 877 seed samples graded "damaged" from all major peanut producing areas of South Africa, 73% were infected.     

Morphology of Schmidt-Lanterman incisures of human shoulder plexus nerves in ontogenesis

The Schmidt-Lanterman incisures are formed in the phase of "loose" myelin at 15-17 weeks of prenatal development with the formation of nodes of Ranvier. The formation of mature Schmidt-Lanterman incisure is completed in the second foetis period. Schmidt-Lanterman incisures distribution in nerve fibres of young men depends on fibre diameter. In the nerves of elderly people the distribution of the incisures in the internode doesn't depend on a fibre diameter. In the nerves of the elderly men the number of incisures is twice as much as that in the young people. The incisural ultrastructure in human nerve is similar to its structure in other mammals.     

Three-dimensional structure of the ciliate Didinium nasutum nucleoli

The nucleolar organization in ciliate Didinium nasutum somatic interphase nuclei was studied using serial ultrathin sections and compared for various physiological states of the cell, namely, fed ciliates, starved ciliates, and dormant cysts. It has been shown that the interphase nucleoli are large structures with a complex architecture: the fibrillar component forms an intricate network in the macronucleus space, while the granular component is located inside this network. The structures looking as individual nucleoli in single sections are actually parts of branched nucleolar networks. The intricate nucleolar networks do not disintegrate after a 30-h starvation; however, the granular component becomes denser and develops numerous cavities filled with fine fibrils of a nonribonucleoprotein nature. In fed D. nasutum, the fibrillar structures on the periphery of nucleoli contain numerous pores (virtually absent in starved cell nucleoli), which can potentially serve for transporting newly synthesized rRNP. Branched nucleolar networks are undetectable in cysts. Their nucleoli are individual structures consisting mainly of the fibrogranular component.     

Description of Paktylenchus tuberosus gen. n., sp. n. (Nematoda: Tylenchinae)

Paktylenchus tuberosus gen.n., sp.n. is described and illustrated from soil around Solanum tuberosum from Kaghan Valley, Pakistan. Paktylenchus gen.n, can be distinguished from the most closely related genus, Agelenchus Andrassy, 1954, by the distinct longitudinal striae, three incisures in the lateral field, and absence of males. Paktylenchus tuberosus shows affinities to Coslenchus Siddiqi, 1978 but differs in having an oblique vagina-to-body axis and a prominent sunken vulva with vulval flaps. Four genera of the subfamily Tylenchinae are also discussed herein.     

Arcyria monticola from Hokkaido, northern Japan: its habit, detailed morphology, and comparison with Arcyria imperialis

Arcyria monticola was found in Hokkaido, northern Japan. This is the first record of its locality, except for the type locality. The linearly expanding capillitium with many coils characterizing A. monticola consists of very elastic longitudinal tubes, short lateral tubes, and many small intricate nets. A longitudinal tube coils around a small intricate net, which forms a core. In contrast, A. imperialis, the other known Arcyria species with a linearly expanding capillitium, has neither short lateral tubes nor nets. The microscopic character of the A. imperialis capillitium is also totally different from that of A. monticola.     

The influence of plane of section on the identification of bone tissue types in amniotes with implications for paleophysiological inferences

The proportion of woven bone (WB) to parallel-fibered bone has been extensively used to infer bone growth rates and resting metabolic rates of extinct organisms. The aim of this study is to test in a variety of amniotes how reliably WB content can be measured using transverse sections. For this, we analyzed femoral transverse mid-diaphyseal thin sections of 14 extant and extinct taxa and the corresponding longitudinal sections for comparative purposes. We used the following characters to identify WB in transverse sections because they are known to be distinct from those observed in parallel-fibered bone: an isotropic bone matrix at tissue scale; an anisotropic microlamellar arrangement in former osteoblast secretory territories at cellular scale; no alignment between osteocytes; and canaliculi running radially from large irregular osteocyte lacunae. Our null hypothesis predicts no differences between the amount of WB quantified in the transverse and longitudinal sections of a given long bone. Qualitatively, when a stripe or a patch of WB was identified in a transverse section, the corresponding stripe or patch of WB was always found at the same location in the corresponding longitudinal section. Quantitatively, a Wilcoxon signed-rank nonparametric paired test did not detect a significant difference in the WB content of the two section planes. Thus, the null hypothesis is not rejected. Considering that paleohistology is a destructive method, we recommend a workflow to efficiently establishing the proportion of WB: quantifying it in transverse sections; preparing and analyzing longitudinal sections only in cases where an ambiguity remains; reanalyzing the corresponding transverse sections.     

The fine structure of the pigment epithelium and photoreceptor cells of the newt,Triturus viridescens dorsalis (Rafinesque)

The morphology of the retinal pigment epithelium and photoreceptor cells has been studied in the common newt Triturus viridescens dorsalis by light, conventional transmission and scanning electron microscopy. The pigment epithelium is formed by a single layer of low rectangular cells, separated by a multilayered membrane (Bruch's membrane) from the vessels of the choriocapillaris. The scleral border of the pigment epithelium is highly infolded and each epithelial cell contains smooth endoplasmic reticulum, myeloid bodies, mitochondria, lysosomes, phagosomes and an oval nucleus. Inner, pigment laden, epithelial processes surround the photoreceptor outer and inner segments. The three retinal photoreceptor types, rods, single cones and double cones, differ in both external and internal appearance. The newt, rod, outer segments appear denser than the cones in both light and electron micrographs, due to a greater number of rod lamellae per unit distance of outer segment and to the presence of electron dense intralamellar bands. The rod outer segments possess deep incisures in the lamellae while the cone lamellae lack incisures. Both rod and cone outer segments are supported by a peripheral array of dendritic processes containing longitudinal filaments which originate in the inner segment. The inner segment mitochondria, forming the rod ellipsoid, arelong and narrow while those in the cone are spherical to oval in shape. The inner segments of all three receptor cell types also contain a glycogen-filled paraboloid and a myoid region, just outside the nucleus, rich in both rough and smooth endoplasmic reticulum. The elongate, cylindrical nuclei differ in density. The rod nuclei are denser than those of the cones, contain clumped chromatin and usually extend further vitreally. Similarly, the cytoplasm of the rod synaptic terminal is denser than its cone counterpart and contains synaptic vesicles almost twice as large as those of the cones. Photoreceptor synapses in rods and cones are established by both superficial and invaginated contacts with bipolar or horizontal cells.     

Mucopolysaccharides in the dog spinal cord and dorsal root ganglia. A histochemical study.

A histochemical study of mucopolysaccharides in the dog spinal cord and dorsal root ganglia is reported in this paper. The histochemical techniques used were the following: PAS, colloidal iron, toluidine blue (pH 5.4 and 3.5), thionine (pH 5.4 and 3.5) and alcian blue 8GX (pH 1 and 2.5). Some histological stains were used also. Two types of neurons could be observed in spinal cord sections stained with colloidal iron techniques. In some neuron a border line of mucosubstances could be seen. In the dorsal root ganglia, different patterns of Nissl bodies distribution in neurons were described. This different distribution of Nissl bodies is associated with different metachromatic colorations of neurons. By using the colloidal iron method, two types of neurons were also revealed in the dorsal root ganglia: some neurons are of a yellow, small-sized and star-shaped type and others are of a light green, larger and round-shaped type. Mucosubstances in the endoneurium and perineurium of nerve fibers, in the Ranvier nodules and in the Schmidt-Lantermann incisures were observed. The possibility that the functional rhythm in some cases might be responsible for the difference in coloration between the dorsal root ganglia neurons is suggested.     

Excitatory glutamate is essential for development and maintenance of the piloneural mechanoreceptor

The piloneural collar in mammalian hairy skin comprises an intricate pattern of circumferential and longitudinal sensory afferents that innervate primary and secondary pelage hairs. The longitudinal afferents tightly associate with terminal Schwann cell processes to form encapsulated lanceolate nerve endings of rapidly adapting mechanoreceptors. The molecular basis for piloneural development, maintenance and function is poorly understood. Here, we show that Nefh-expressing glutamatergic neurons represent a major population of longitudinal and circumferential sensory afferents innervating the piloneural collar. Our findings using a VGLUT2 conditional-null mouse model indicate that glutamate is essential for innervation, patterning and differentiation of NMDAR(+) terminal Schwann cells during piloneural collar development. Similarly, treatment of adult mice with a selective NMDAR antagonist severely perturbed piloneural collar structure and reduced excitability of these mechanosensory neurons. Collectively, these results show that DRG-derived glutamate is essential for the proper development, maintenance and sensory function of the piloneural mechanoreceptor.     

Calmodulin immunolocalization in outer segments of Xenopus laevis photoreceptors

Because adaptation of vertebrate photoreceptors to light is mediated by changes in the level of calcium in their outer segments (OS), proteins that bind calcium are important in phototransduction. This study has used immunofluorescence to investigate the distribution of the calcium-binding protein calmodulin within photoreceptor OS dissociated from amphibian ( Xenopus laevis) retinas. The OS of rods and cones had a streak of fluorescence to calmodulin at the ciliary axoneme. The OS of rods (but not cones) also displayed regularly spaced puncta of anti-calmodulin fluorescence along longitudinal lines coinciding with their multiple incisures. This location of calmodulin immunofluorescence closely matches the known location of microtubules within the OS of amphibian rods and cones. These findings provide evidence that calmodulin is closely associated with the microtubules of both the axonemal and incisural cytoskeletal systems in OS, and suggest that this association is important for calmodulin function in photoreceptors.     

Fine structural and immunohistochemical studies of goat adenohypophysial cells

Summary The fine structure of each type of anterior pituitary cell in the male goat was studied through the application of a superimposition technique in which adjacent thick sections were used to identify individual cells beforehand by light-microscopic immunohistochemistry. A cone of the pars intermedia protrudes into the pars anterior, being surrounded by the narrow pituitary cleft; the immunohistochemical appearances of the cells forming the cone resemble those of the pars anterior. Several follicles appear in the pars anterior. Ultrastructurally GH cells resemble prolactin cells. The secretory granules of both types are spherical; the diameter of the former is about 340 nm, whereas that of the latter is about 440 nm. ACTH cells are polygonal in shape with secretory granules, about 180 nm in diameter, scattered throughout the cytoplasm. TSH cells, which are spherical in shape, contain the smallest secretory granules, 150 nm in diameter. The highly electron-dense LH cells contain numerous secretory granules about 210 nm in diameter. Their nuclei are irregular with incisures. Thus, the anterior pituitary cells of the goat are ultrastructurally characteristic and species-specific.