The bending properties of single osteons

The bending properties of two fully calcified osteon types (longitudinal and alternate) have been investigated in 62 cylindrical samples by applying the technique of three-point bending loading. The bending of each sample was recorded using a microwave micrometer based on the cavity and pulse technique. It has been shown that alternate osteons are better able to withstand bending stress than longitudinal ones. This result offers a definitive explanation for the high concentration of transverse lamellae in pathologically bowed bone shaft.     

The anisotropic material constitutive models for the human cornea

This paper presents an anisotropic analysis model for the human cornea. The model is based on the assumption that the fibrils in the cornea are organised into lamellae, which may have preferential orientation along the superior-inferior and nasal-temporal directions, while the alignment of lamellae with different orientations is assumed to be random. Hence, the cornea can be regarded as a laminated composite shell. The constitutive equation describing the relationships between membrane forces, bending moments, and membrane strains, bending curvatures are derived. The influences of lamella orientations and the random alignment of lamellae on the stiffness coefficients of the constitutive equation are discussed.     

金鱼卵母细胞发育过程中环孔片层的电镜观察

本文研究了环孔片层在金鱼卵母细胞发育过程中的一些形态变化。描述卵母细胞发育的五个阶段中每个阶段环礼片层的分布以及排列的情况。结果表明随着卵母细胞的生长,环孔片层的出现率和每束中片层的数目都逐渐增加。除了在卵原细胞以外,环礼片层一般都分布在远离细胞核的卵质中。尚观察到孔结合物质和片层间物质与多核蛋白体有联系。我们从观察结果推测金鱼卵母细胞的环孔片层可能是由粗面内质网衍生而成。     

Distribution of osteonic and interstitial components in the human femoral shaft with reference to structure, calcification and mechanical properties

This paper analyzes the distribution of osteons and interstitial bone in the femoral compacta according to their structure, degree of calcification and mechanical properties. Three cross sections, 100 microns thick, each located 1 cm from the next, were prepared by grinding from the middle third of a human femoral shaft. Starting from the premise that, in lamellar bone, lamellae whose fiber bundles and crystallites have a longitudinal course withstand loading by tension, whereas those whose fiber bundles and crystallites have a transversal course withstand loading by compression, each osteon and fragment of interstitial bone has been given a number recording the percentage of its surface consisting of lamellae with transversally oriented fiber bundles and crystallites (bright under the polarizing microscope). The degree of calcification of the same structures was determined micro-radiographically. The distribution of both osteons and interstitial bone was assessed using a tungsten grid for reference. The total surface of each bone microstructure, and the percentage of that surface consisting of bright lamellae, were all calculated using a Zeiss Video-plan. Our results confirm the view that the distribution of both osteons and interstitial bone is mainly related to their structure--and hence to their mechanical properties. In addition, bone remodeling seems to be most active in areas capable of supporting tensile stress.     

Scanning electron microscopy of the gills of a freshwater catfish,Rita Rita

Variations in the gross morphology and surface architecture of the gill filaments and secondary lamellae of a freshwater catfish (Rita rita) have been investigated using scanning electron microscopy. Heterogeneity of the gill has been correlated with the distribution of lamellar water-flow at different regions of a gill filament. Higher lamellar water flow (cc/pore/cmH₂O/sec) was estimated for the middle region of the filaments. The filaments are covered with epithelial cells whose surface is provided with well-developed microridges. The lamellae are generally covered with microvillous epithelial cells. The variations in surface architecture of the gill filaments and secondary lamellae have been correlated with their probable functions.     

Collagen fibre orientation in the annulus fibrosus of intervertebral disc during bending and torsion measured by x-ray diffraction

X-ray diffraction has been used to measure the orientation of the collagen fibres in the ventral annulus fibrosus of intact L1/2 rabbit intervertebral disc during in vitro bending and torsion. Fibres are tilted with respect to the axis of the spine. As predicted by theory, fibre tilt decreases in those regions of the annulus which are stretched by bending but increases in the slackened regions. Good agreement with the quantitative predictions of bending theory was obtained in three of the six series of experiments, the predicted trend being found in all six. Tilt direction alternates in successive lamellae of the annulus. When discs were subjected to both clockwise and anticlockwise torsion of 5°, the two families of titled fibres reoriented in the expected directions.     

Microstructural mechanical study of a transverse osteon under compressive loading: The role of fiber reinforcement and explanation of some geometrical and mechanical microscopic properties

This Finite Element study aims at understanding the transverse osteon as a composite microstructure, and at differentiating the actions of each of its main components and their interactions. Three components of the osteon have been distinguished: the lamellae mineral-collagen matrix, the lamellae mineral-collagen reinforcement fibers and the Haversian canal content made of intracortical fluid and soft tissues. Numerical compression experiments have been performed, varying the microstructure properties. Our results show that fiber reinforcement of transverse osteons is only efficient at resisting dynamic compressive loadings, but that the improvement of the static compressive properties is very poor. Furthermore, the modeled stress distribution within the matrix and reinforcement fibers may explain why transverse osteons are often limited to a small number of lamellae (<8) and why internal lamellae could be stiffer than external ones.     

THE FINE STRUCTURE OF THE CILIA FROM CTENOPHORE SWIMMING-PLATES

The ctenophore swimming-plate has been examined with the electron microscope. It has been recognized as an association of long cilia in tight hexagonal packing. One of the directions of the hexagonal packing is parallel to the long edge of the swimming-plate and is perpendicular to the direction of the ciliary beat. All the cilia in the swimming-plate are identically oriented. The effective beat in the movement of the swimming-plate is directed towards the aboral pole of the animal, and this is also the side of the unpaired peripheral filament in all the cilia. The direction of the ciliary beat is fixed in relation to the position of the filaments of the cilia. The swimming-plate cilium differs from other types of cilia and flagella in having a filament arrangement that can be described as 9 + 3 as opposed to the conventional 9 + 2 pattern. The central filaments appear in a group of two "tubular" filaments and an associated compact filament. The compact filament might have a supporting function. It has been called "midfilament." Two of the peripheral nine filaments (Fig. 1, Nos. 3 and 8) are joined to the ciliary membrane by means of slender lamellae, which divide the cilium into two unequal compartments. These lamellae have been called "compartmenting lamellae." Some observations of the arrangement of the compartmenting lamelae indicate that they function by cementing the cilia together in lateral rows. The cilia of the rows meet at a short distance from each other, leaving a gap of 30 A only. The meeting points are close to the termini of the compartmenting ridges. An electron-dense substance is sometimes seen bridging the gap. Some irregularities are noted with regard to the arrangement of the compartmenting lamellae particularly at the peripheral rows of cilia. In many cilia in these rows there are small vesicles beneath the ciliary membrane.     

Filipin-sterol complexes in disrupted myelin in the rat

Using filipin as a cytochemical probe for cholesterol we have compared the distribution of filipin labelling in mildly disrupted myelin and normal myelin. The myelin lamellae in rat sciatic nerve were separated either by hypotonic saline (0.035-0.07 M) or nerve section (24-32 h) before aldehyde fixation and filipin treatment. Myelin separation was assessed in ultrathin sections and filipin distribution in freeze-fracture replicas. In separated myelin lamellae filipin labelling was similar throughout the myelin sheath while in normal control myelin filipin occurred most in the outer (abaxonal), least in the inner (adaxonal) and intermediate in the middle lamellae. It is concluded that this heterogeneous filipin labelling in normal myelin is a result of diffusion gradients to filipin within the myelin sheath and that in vivo cholesterol is uniformly distributed throughout all the lamellae of the myelin sheath. The site of the diffusion barrier to filipin within normal myelin is considered.     

Effects of the basic multicellular unit and lamellar thickness on osteonal fatigue life

A remodeling cycle sets the size of the osteon and associated lamellae in the basic multicellular unit. Treatments and aging affect these micro-structural features. We previously demonstrated decreased fatigue life with an unexplained mechanism and decreased osteon size in cortical bone treated with high-dose bisphosphonate. Here, three finite element models were examined: type-1: a single osteon, as a homogeneous unit and with heterogeneous lamellae and interlamellae, type-2: a control, interstitial-only tissue and type-3: the osteon with cement line, set within the interstitial tissue. Models were loaded in simulated, sinusoidal bending fatigue. As osteon size was decreased, lamellar number and lamellar thickness were incrementally adjusted for each model. As hypothesized, lamellae within the larger type-1 models attained greater cycles to failure and the addition of an osteon to type-2 models (generating a type-3 model set) yielded increased fatigue life. However, as the osteon size was decreased, the potential for compressive damage nucleation was increased within the lamellae of the osteons versus the interstitium. Also, osteons with fewer, thicker lamellae displayed increased fatigue life. Osteonal microstructure plays a role in damage initiation location, especially when BMU size is smaller. Previous findings by us and others could partially be explained by this further understanding of increased probability for damage nucleation in smaller osteons.     

Basophilic Lamellar Systems in the Crayfish Spermatocyte

Histochemical procedures for the demonstration of RNA have shown the presence of intensely basophilic bodies in the cytoplasm of spermatocytes of the crayfish, Cambarus virilis. The staining of thick sections, cut alternately with thin sections for electron microscopy, has permitted identification of the basophilic bodies with two types of lamellar systems. One of these, a set of straight annulate lamellae, is restricted to meiotic prophase. The second type of lamellar systems has been found from late prophase to early spermatid stages. It consists of an ellipsoidal lamellar set which intersects a number of straight lamellae. Within the region of intersection, the ellipsoidal lamellae break up into an array of small tubules of about 150 A diameter. The term tubulate lamellar system was chosen to designate this type of lamellar complex. Small RNA-containing granules could not be detected in annulate lamellar systems. While there are a few granules in the marginal regions of the tubulate lamellar system, their distribution cannot be responsible for the basophilia which is intense within all regions of the lamellar body.     

ELECTRON MICROSCOPY OF THE PACINIAN CORPUSCLE

The Pacinian corpuscle has a framework of cytoplasmic lamellae arranged concentrically in the outer zone, and bilaterally in the core. Between these is an intermediate growth zone. The inner core shows an unexpected complexity in that its component lamellae are arranged in two symmetrical groups of nested cytoplasmic sheets. Longitudinal tissue spaces form clefts separating the two groups. The perikarya of the core lamellae lie in or near the intermediate growth zone, and send arms into the clefts. The arms then branch and terminate as lamellae which interdigitate with those of neighboring cells. The single nerve fiber loses its myelin sheath just before it reaches the inner core but retains its Schwann cell cytoplasmic covering for a short additional distance. The Schwann sheath is not continuous with the lamellae of the inner core. Inside the core the fiber contains a striking circumferential palisade of radially disposed mitochondria. The fiber does not arborize. Vascular capillaries penetrate the hilar region of the corpuscle only as far as the myelinated sheath of the nerve, and they have not been seen elsewhere in the corpuscle. There is direct continuity between the clefts of the core and tissue spaces in the vicinity of the capillaries. It is likely that this provides a route whereby metabolites reach the active nerve ending, as well as the cells of the growth zone. The outer zone consists of at least 30 flattened concentric cytoplasmic lamellae separated from one another by relatively wide fluid-filled spaces. Collagenous fibrils are present, particularly on the outer surface of lamellae, and tend to be oriented circularly. The girdle of proliferating cells constituting the growth zone, which is prominent in corpuscles from young animals, is the layer from which the outer lamellae are derived. Osmotic forces probably elevate the lamellae, and maintain turgor pressure.     

Annulate lamellae in an undifferentiated mouse marrow cell

Annulate lamellae have been observed in undifferentiated cells in the marrow of mice treated with urethan. They were not found in any recognizable hematopoietic element in either urethan- or saline-injected controls. The possibility has been raised that this cytoplasmic organelle is confined to the most primitive marrow cell and may be an identifying feature of 'stem cells'.     

Annulate lamellae in human tumor cells.

Annulate lamellae have been found in a primitive neuroectodermal tumor, a metastatic cerebellar tumor, a testicular seminoma, a retinoblastoma and three melanomas. These annulate lamellae are arranged in stacked parallel arrays in the cytoplasm of tumor cells. The number of annulate lamellae observed to comprise a single stack varies from 2--4 in the seminoma tumor to 5--18 in the cerebellar tumor. Although the functional significance of annulate lamallae is still unknown, in many instances they have been found to be continuous with rough-surfaced cisternae of the endoplasmic reticulum and ribosomes have been demonstrated on the surface of annulate lamellae. This may suggest that annulate lamellae participate in protein synthesis.     

Fractionation of the thylakoid membranes from tobacco. A tentative isolation of 'end membrane' and purified 'stroma lamellae' membranes

Thylakoids isolated from tobacco were fragmented by sonication and the vesicles so obtained were separated by partitioning in aqueous polymer two-phase systems. By this procedure, grana vesicles were separated from stroma exposed membrane vesicles. The latter vesicles could be further fractionated by countercurrent distribution, with dextran-polyethylene glycol phase systems, and divided into two main populations, tentatively named 'stroma lamellae' and 'end membrane'. Both these vesicle preparations have high chlorophyll a/b ratio, high photosystem (PS) I and low PS II content, suggesting their origin from stroma exposed regions of the thylakoid. The two vesicle populations have been compared with respect to biochemical composition and photosynthetic activity. The 'end membrane' has a higher chlorophyll a/b ratio (5.7 vs. 4.7), higher P700 content (4.7 vs. 3.3 mmol/mol of chlorophyll). The 'end membrane' has the lowest PS II content, the ratio PS I/PS II being more than 10, as shown by EPR measurements. The PS II in both fractions is of the beta-type. The decay of fluorescence is different for the two populations, the 'stroma lamellae' showing a very slow decay even in the presence of K3Fe(CN)6 as an acceptor. The two vesicle populations have very different surface properties: the end membranes prefer the upper phase much more than the stroma lamellae, a fact which was utilized for their separation. Arguments are presented which support the suggestion that the two vesicle populations originate from the grana end membranes and the stroma lamellae, respectively.     

The micromechanics versus the macromechanics of cortical bone--a comprehensive presentation

Secondary cortical bone is a complicated patchwork of structures which can be viewed as a hierarchy of four different orders. As far as the biomechanical properties of cortical bone are concerned, the lamellae is the most important of the four. The relative distribution of longitudinal lamellae (whose fiber bundles and crystallites have a longitudinal course and withstand loading by tension) with respect to transverse lamellae (whose fiber bundles and crystallites have a transverse course and withstand loading by compression) governs the mechanical properties of bone at macroscopic level both in normal and pathological conditions.     

A multilamellate body in the granulated hypophysial cells of the teleost,Hemihaplochromis philander

Summary A multilamellate body (MLB), bearing close resemblance to an array of annulate lamellae, has been observed in several adenohypophysial cell types of the teleost, Hemihaplochromis philander. In longitudinal section, each MLB comprises a ladder-like row of 12–50 sausage-shaped profiles, termed lamellae. A few lamellae in each section show connections with the endoplasmic reticulum. Apposition of paired lamellar membranes at regularly spaced intervals results in a beaded appearance, whereby sites of membrane apposition are probable pore sites. The MLB differs from annulate lamellae in having poorly developed pores and closer packing of lamellae. It is suggested that the MLBs described here, may represent annulate lamellae at a stage of development or break-down when pores are incomplete.Acknowledgement is made to the South African Council for Scientific and Industrial Research for financial aid towards this work. The authors also wish to thank the Natal Parks, Game and Fish Preservation Board for providing specimens     

Threat-Sensitive Responses to Predator Attacks in a Damselfly

The threat sensitivity hypothesis predicts that prey species assess and adjust their behavior flexibly in accordance with the magnitude of the threat imposed by a predator. We tested this hypothesis with regard to escape behavior and thanatosis (feigning of death to escape predation) in larvae of the damselfly Ischnura elegans. We manipulated the perceived predation threat of the larvae by changing three factors: lamellae autotomy (an escape strategy where animals sacrifice a body part when grasped by a predator; lamellae present or absent), kairomone type (odors released by predators; control, dragonfly kairomones or fish kairomones), and population of origin (fishpond or fishless pond). We demonstrated that thanatosis increased survival both when confronted with dragonfly and fish predators. We could show, for the first time, costs of past autotomy to be predator‐dependent: larvae without lamellae suffered higher predation mortality but only in the presence of a dragonfly predator and not in the presence of a fish predator. This is in accordance with the observed reduced escape speed of larvae after autotomy, which may affect escape probability toward dragonfly predators but not to the very fast fish predators. Unexpectedly, kairomone type did not affect the escape response of the larvae. In accordance with the threat sensitivity hypothesis, after an unsuccessful attack, larvae without lamellae had a higher frequency to enter thanatosis than larvae with lamellae and larvae from the fishpond showed longer thanatosis durations than larvae from the fishless pond. Consistent with the hypothesis, the reaction of the larvae to a simulated attack depended jointly on lamellae status and population. In fishless ponds, larvae with lamellae swam away more frequently than larvae without lamellae; in fishponds both groups almost never swam away and relied mostly upon immobility. Given the obvious benefits of adaptively varying escape responses we hypothesize this threat sensitivity to be widespread. Moreover, we argue that former inconsistencies between studies with regard to escape behavior may have been partly because of such adaptive variation.     

Orientation of collagen fibers at the boundary between two successive osteonic lamellae and its mechanical interpretation

By applying an original technique, an investigation has been carried out to determine the orientation of collagen fibrils at the boundary between two successive lamellae in alternate osteons. Evidence is reported that the predominant fiber direction does not change abruptly from one lamella to the next; there is an intermediate system of criss-crossed fibers whose main orientation makes an angle of nearly 45 degrees with the direction of the fibers in the two adjacent lamellae. Taking a composite orthogonally reinforced laminate as a model, a mechanical interpretation of this intermediate system of collagen fibers is given.     

Relationships between annulate lamellae and filament bundles in oocytes of the zebrafish,Brachydanio rerio

Summary Bundles of filaments have been observed in the vitellogenic oocyte of the zebrafish, Brachydanio rerio; and these filaments illustrate a close spatial and structural relationship to annulate lamellae. The filaments range from 6–8 nm in diameter, and the annulate lamellae may cap both rounded ends of the bundle as well as extend parallel to the surface of the filament bundles. The ends of the filaments can be observed to exhibit an apparent termination in close relation to pore margins of the annulate lamellae, the membrane of the interpore regions of the annulate lamellae, as well as many nearby polyribosomes. The possible functional significance of this unique relationship is discussed in reference to a recent hypothesis regarding the function of annulate lamellae.