Locomotory aspects of squid mantle structure

Morphological aspects of squid ( Loligo, Lolliguncula ) mantle relevant to locomotory function were studied. Methods used included polarized light microscopy of frozen sections of untreated tissue taken from animals immediately after death and electron microscopy.
The mantle consists of circular and radial muscles arranged in alternating rings along the whole length of the mantle. The muscle is obliquely striated. Connective tissue fibres are found in the body of the muscle and in the outer and inner tunics. The outer tunic consists of layers of large collagenous fibres. The fibres run in superimposed right- and left-handed helical courses that lie at an angle of 27° to the long axis of the animal. The tunics and the intramuscular connective fibres are thought to resist length changes in the mantle while permitting the changes in girth required for the jet power stroke. Both the intramuscular and the tunic fibre systems may provide elastic energy for the return phase of the jet cycle. Tunic fibres appear to be a geodesic tensile reinforcing system ensuring smooth shape changes in the mantle.     

Opening angle and residual strain in a three-layered model of pig oesophagus

Studies of various biological tissues have shown that residual strains are important for tissue function. Since a force balance exists in whole wall thickness specimens cut radially, it is evident that layer separation is an important procedure in the understanding of the meaning of residual stresses and strains. The present study investigated the zero-stress state and residual strain distribution in a three-layer model of the pig oesophagus. The middle part of the oesophagus was obtained from six slaughterhouse pigs. Four 3-mm-wide rings were serially cut from each oesophagus. Two of them were used for separating the wall into mucosa-submucosa, inner and outer muscle layers. The remaining two rings were kept as intact rings. The inner and outer circumferences and wall thickness of different layers in intact and separated rings were measured from the digital images in the no-load state and zero-stress state. The opening angle was measured and the residual strain at the inner and outer surface of different layers and the intact wall were computed. Compared with intact sectors (62.8+/-9.8 degrees ), the opening angles were smaller in the inner muscle sectors (37.2+/-11.4 degrees , P<0.01), whereas the opening angles of mucosa-submucosa (63.9+/-6.8 degrees ) and outer muscle sectors (63.9+/-6.8 degrees ) did not differ (P>0.1). Referenced to the zero-stress state of the intact sectors, the inner and outer residual strains of the intact rings was -0.128+/-0.043 and outer residual strain was 0.308+/-0.032. Referenced to the "true" zero-stress state of separated three-layered sectors, the inner residual strain of intact rings were -0.223+/-0.021 (P<0.01) and 0.071+/-0.022 (P<0.01). Referenced to the "true" zero-stress state, the residual strain distribution of different layers in intact rings was shown that the inner surface residual strain was negative at mucosa-submucosa and inner muscle layers and was positive at outer muscle layer, whereas the outer surface residual strain was negative at the mucosa-submucosa layer and positive at the inner and outer muscle layers. For the separated different layered rings, the inner residual strain was negative and outer residual strain was positive; however, the absolute values did not differ (P>0.1). In conclusion, it is possible to microsurgically separate the oesophagus into three layers, i.e., mucosa-submucosa, inner muscle and outer muscle layers, the residual strain differ between the layers, and the residual strain distribution was more uniform after the layers were separated.     

Evidence for a radial strain gradient in apple fruit cuticles

Main conclusion

The morphological outer side of the apple fruit cuticle is markedly more strained than the inner side. This strain is released upon wax extraction. This paper investigates the effect of ablating outer and inner surfaces of isolated cuticular membranes (CM) of mature apple (Malus × domestica) fruit using cold atmospheric pressure plasma (CAPP) on the release of strain after extraction of waxes. Strain release was quantified as the decrease in area of CM discs following CAPP treatment and subsequent solvent extraction of wax. Increasing duration of CAPP treatment proportionally decreased CM mass per unit area. There was no difference in mass loss rate between CAPP treatments of outer or inner surfaces. Also, there was no difference in surface area of CMs before and after CAPP treatment. However, upon subsequent wax extraction, surface area of CMs decreased indicating the release of strain. Increasing the duration of CAPP treatment resulted in increasing strain release up to 47.7 ± 8.0 % at 20 min when CAPP was applied to the inner surface. In contrast, strain release was independent of CAPP duration averaging about 12.1 ± 0.6 % when applied to the outer surface of the CM. Our results provide evidence for a marked gradient of strain between the outer side (strained) and the inner side of the CM (not strained) of mature apple fruit.     

Ontogeny of squid mantle function: changes in the mechanics of escape-jet locomotion in the oval squid,Sepioteuthis lessoniana lesson, 1830

In Sepioteuthis lessoniana, the oval squid, ontogenetic changes in the kinematics of the mantle during escape-jet locomotion imply a decline in the relative mass flux of the escape jet and may affect the peak weight-specific thrust of the escape jet. To examine the relationship between ontogenetic changes in the kinematics of the mantle and the thrust generated during the escape jet, we simultaneously measured the peak thrust and the kinematics of the mantle of squid tethered to a force transducer. We tested an ontogenetic series of S. lessoniana that ranged in size from 5 to 40 mm dorsal mantle length (DML). In newly hatched squids, thrust peaked 40 ms after the start of the escape jet and reached a maximum of between 0.10 mN and 0.80 mN. In the largest animals, thrust peaked 70 ms after the start of the escape jet and reached a maximum of between 18 mN and 110 mN. Peak thrust was normalized by the wet weight of the squid and also by the cross-sectional area of the circumferential muscle that provides power for the escape jet. The weight-specific peak thrust of the escape jet averaged 0.36 in newly hatched squid and increased significantly to an average of 1.5 in the largest squids measured (P < 0.01). The thrust per unit area of circumferential muscle averaged 0.25 mN/mm(2) in hatchlings and increased significantly to an average of 1.4 mN/mm(2) in the largest animals tested (P < 0.01). The impulse of the escape jet was also lowest in newly hatched individuals (1.3 mN. s) and increased significantly to 1000 mN. s in the largest squids measured (P < 0.01). These ontogenetic changes in the mechanics of the escape jet suggest (1) that propulsion efficiency of the exhalant phase of the jet is highest in hatchlings, and (2) that the mechanics of the circumferential muscles of the mantle change during growth.     

Regional epicardial strain in the embryonic chick heart during the early looping stages

Epicardial strains were measured in Hamburger-Hamilton stage 11 and 12 embryonic chick hearts (1.6-2.0 days of incubation). These stages include part of the early phase of cardiac looping, as the initially straight heart tube bends and twists to form a curved c-shaped tube. By analyzing the motion of microbeads placed on the myocardial surface, we measured strains near the outer curvature, in the central region, and near the inner curvature of the primitive ventricle. No significant differences in strain were found between stages. Relative to end diastole, all three regions shortened by about 10% during systole in the circumferential direction, and the outer curvature shortened longitudinally by about 5%. In contrast, and unlike strains in older hearts, the inner curvature and central regions elongated by approximately 5-10% in the longitudinal direction during systole. These results are consistent with microstructural data and suggest that the material properties of the outer curvature are relatively isotropic, whereas the properties of the central and inner curvature regions are orthotropic, with contractile stress exerted primarily in the circumferential direction.     

Complex distributions of residual stress and strain in the mouse left ventricle: experimental and theoretical models

 Most soft biological tissues, including ventricular myocardium, are not stress free when all external loads are removed. Residual stress has implications for mechanical performance of the heart, and may be an indicator of patterns of regional growth and remodeling. Cross-sectional rings of arrested ventricles opened up when a radial cut was made (initial mean opening angles were 64 ± 17°), but further circumferential cuts revealed the presence of additional residual stresses in the tissue with further opening of the rings. In normal mouse hearts, the inner half of a short-axis ring opened more than the outer half, and this change was dependent on apex–base location. At the apex the inner section vs. outer section opening angles were 226 ± 47° vs. 89 ± 28°, while at the base the same two angles were 160 ± 30° vs. 123 ± 35°. A simple theoretical cylindrical shell model with incompressible hyperelastic material properties was used to model the experimental deformations based on the cutting experiments. The model predicts different residual stress fields depending on the nature of the opening after the circumferential cut (which is done after the conventional radial cut). The observed opening angles were consistent with steep stress gradients near the endocardium compared with those predicted if the first cut was assumed to relieve all residual stresses. These results imply a more complex distribution of residual stress and strain in ventricular myocardium than previously thought. Received: 23 May 2002 / Accepted: 30 September 2002 We would like to acknowledge the surgical skills and data analysis of Zuangjie Li. This work was supported in part by National Heart, Lung, and Blood Institute Grants HL-43026 and HL-64321.     

A new model for periostracum and shell formation in Unionidae (Bivalvia, Mollusca)

The periostracum in Unionidae consists of two layers. The outer one is secreted within the periostracal groove, while the inner layer is secreted by the epithelium of the outer mantle fold. The periostracum reaches its maximum thickness at the shell edge, where it reflects onto the shell surface. Biomineralization begins within the inner periostracum as fibrous spheruliths, which grow towards the shell interior, coalesce and compete mutually, originating the aragonitic outer prismatic shell layer. Prisms are fibrous polycrystalline aggregates. Internal growth lines indicate that their growth front is limited by the mantle surface. Transition to nacre is gradual. The first nacreous tablets grow by epitaxy onto the distal ends of prism fibres. Later growth proceeds onto previously deposited tablets. Our model involves two alternative stages. During active shell secretion, the mantle edge extends to fill the extrapallial space and the periostracal conveyor belt switches on, with the consequential secretion of periostracum and shell. During periods of inactivity, only the outer periostracum is secreted; this forms folds at the exit of the periostracal groove, leaving high-rank growth lines. Layers of inner periostracum are added occasionally to the shell interior during prolonged periods of inactivity in which the mantle is retracted.     

On the absence of circular muscle elements in the body wall of Dysponetus pygmaeus (Chrysopetalidae, 'Polychaeta', Annelida)

The annelid body wall generally comprises an outer layer of circular muscle fibres and an inner layer of longitudinal muscle fibres as well as parapodial and chaetal muscles. An investigation of Dysponetuspygmaeus (Chrysopetalidae) with confocal laser scanning microscopy showed that circular muscles are entirely absent. Further studies indicate that this feature is characteristic for all Chrysopetalidae. A scrutiny of the literature showed a similar situation in many other polychaetes. This lack of circular muscle fibres may either be due to convergence or represent a plesiomorphic character. Since circular muscles are very likely important for burrowing forms but not necessary for animals which proceed by movements of their parapodial appendages or cilia, this problem is also related to the question of whether the ancestral polychaete was epi‐ or endobenthic.     

Three-dimensional reconstruction of the musculature of Cossura pygodactylata Jones, 1956 (Annelida: Cossuridae)

The musculature of adult specimens of Cossura pygodactylata was studied by means of F-actin labelling and confocal laser scanning microscopy (CLSM). Their body wall is comprised of five longitudinal muscle bands: two dorsal, two ventral and one ventromedial. Complete circular fibres are found only in the abdominal region, and they are developed only on the border of the segments. Thoracic and posterior body regions contain only transverse fibres ending near the ventral longitudinal bands. Almost-complete rings of transverse muscles, with gaps on the dorsal and ventral sides, surround the terminal part of the pygidium. Four longitudinal bands go to the middle of the prostomium and 5–14 paired dorso-ventral muscle fibres arise in its distal part. Each buccal tentacle contains one thick and two thin longitudinal muscle filaments; thick muscle fibres from all tentacles merge, forming left and right tentacle protractors rooted in the dorsal longitudinal bands of the body wall. The circumbuccal complex includes well-developed upper and lower lips. These lips contain an outer layer of transverse fibres, and the lower lip also contains inner oblique muscles going to the dorsal longitudinal bands. The branchial filament contains two longitudinal muscle fibres that do not connect with the body musculature. The parapodial complex includes strong intersegmental and segmental oblique muscles in the thoracic region only; chaetal retractors, protractors and muscles of the body wall are present in all body regions. Muscle fibres are developed in the dorsal and ventral mesenteries. One semi-circular fibre is developed on the border of each segment and is most likely embedded in the dissepiment. The intestine has thin circular fibres along its full length. The dorsal blood vessel has strong muscle fibres that cover its anterior part, which is called the heart. It consists of short longitudinal elements forming regular rings and inner partitions. The musculature of C. pygodactylata includes some elements that are homologous with similar muscular components in other polychaetes (i.e., the body wall and most parapodial muscles) and several unique features, mostly at the anterior end.     

Role of aerobic and anaerobic circular mantle muscle fibers in swimming squid: electromyography

Circular mantle muscle of squids and cuttlefishes consists of distinct zones of aerobic and anaerobic muscle fibers that are thought to have functional roles analogous to red and white muscle in fishes. To test predictions of the functional role of the circular muscle zones during swimming, electromyograms (EMGs) in conjunction with video footage were recorded from brief squid Lolliguncula brevis (5.0-6.8 cm dorsal mantle length, 10.9-18.3 g) swimming in a flume at speeds of 3-27 cm s(-1). In one set of experiments, in which EMGs were recorded from electrodes intersecting both the central anaerobic and peripheral aerobic circular mantle muscles, electrical activity was detected during each mantle contraction at all swimming speeds, and the amplitude and frequency of responses increased with speed. In another set of experiments, in which EMGs were recorded from electrodes placed in the central anaerobic circular muscle fibers alone, electrical activity was not detected during mantle contraction until speeds of about 15 cm s(-1), when EMG activity was sporadic. At speeds greater than 15 cm s(-1), the frequency of central circular muscle activity subsequently increased with swimming speed until maximum speeds of 21-27 cm s(-1), when muscular activity coincided with the majority of mantle contractions. These results indicate that peripheral aerobic circular muscle is used for low, intermediate, and probably high speeds, whereas central anaerobic circular muscle is recruited at intermediate speeds and used progressively more with speed for powerful, unsteady jetting. This is significant because it suggests that there is specialization and efficient use of locomotive muscle in squids.     

Ontogenetic changes in fibrous connective tissue organization in the oval squid, Sepioteuthis lessoniana Lesson, 1830.

Ontogenetic changes in the organization and volume fraction of collagenous connective tissues were examined in the mantle of Sepioteuthis lessoniana, the oval squid. Outer tunic fiber angle (the angle of a tunic collagen fiber relative to the long axis of the squid) decreased from 33.5 degrees in newly hatched animals to 17.7 degrees in the largest animals studied. The arrangement of intramuscular collagen fiber systems 1 (IM-1) and 2 (IM-2) also changed significantly during ontogeny. Because of the oblique trajectory of the IM-1 collagen fibers, two fiber angles were needed to describe their organization: (1) IM-1(SAG), the angle of an IM-1 collagen fiber relative to the squid's long axis when viewed from a sagittal plane and (2) IM-1(TAN), the angle of an IM-1 collagen fiber relative to the squid's long axis when viewed from a plane tangential to the outer curvature of the mantle. The sagittal component (IM-1(SAG)) of the IM-1 collagen fiber angle was lowest in hatchling squid (32.7 degrees ) and increased exponentially during growth to 43 degrees in squid with a dorsal mantle length (DML) of 15 mm. In squid larger than 15 mm DML, IM-1(SAG) fiber angle did not change. The tangential component (IM-1(TAN)) of IM-1 collagen fiber angle was highest in hatchling squid (39 degrees ) and decreased to 32 degrees in the largest squid examined. IM-2 collagen fiber angle (the angle of an IM-2 collagen fiber relative to the outer surface of the mantle) was lowest in hatchling squid (34.6 degrees ) and increased exponentially to about 50 degrees in 15-mm DML animals. In squid larger than 15 mm DML, IM-2 fiber angle increased slightly with size. The volume fraction of collagen in IM-1 and IM-2 increased 68 and 36 times, respectively, during growth. The ontogenetic changes in the organization of collagen fibers in the outer tunic, IM-1, and IM-2 may lead to ontogenetic differences in the kinematics of mantle movement and in elastic energy storage during jet locomotion.     

Nitric oxide synthase immunoreactivity in the enteric nervous system of the developing human digestive tract

We have investigated indirectly the presence of nitric oxide in the enteric nervous system of the digestive tract of human fetuses and newborns by nitric oxide synthase (NOS) immunocytochemistry and nicotinamide adenine dinucleotide phosphate diaphorase (NADPHd) histochemistry. In the stomach, NOS immunoactivity was confined to the myenteric plexus and nerve fibres in the outer smooth musculature; few immunoreactive nerve cell bodies were found in ganglia of the outer submucous plexus. In the pyloric region, a few nitrergic perikarya were seen in the inner submucous plexus and some immunoreactive fibres were found in the muscularis mucosae. In the small intestine, nitrergic neurons clustered just underneath or above the topographical plane formed by the primary nerve strands of the myenteric plexus up to the 26th week of gestation, after which stage, they occurred throughout the ganglia. Many of their processes contributed to the dense fine-meshed tertiary nerve network of the myenteric plexus and the circular smooth muscle layer. NOS-immunoreactive fibres directed to the circular smooth muscle layer originated from a few NOS-containing perikarya located in the outer submucous plexus. In the colon, caecum and rectum, labelled nerve cells and fibres were numerous in the myenteric plexus; they were also found in the outer submucous plexus. The circular muscle layer had a much denser NOS-immunoreactive innervation than the longitudinally oriented taenia. The marked morphological differences observed between nitrergic neurons within the developing human gastrointestinal tract, together with the typical innervation pattern in the ganglionic and aganglionic nerve networks, support the existenc of distinct subpopulations of NOS-containing enterice neurons acting as interneurons or (inhibitory) motor neurons.     

Differentiation and Growth of the Brachiopod Mantle

Cell differentiation in the mantle edge of Notosaria, Thecidelhnaand Glottidia, representing respectively, the impunctate andpunctate calcareous articulate and chitinophosphatic inarticulatebrachiopods, is described. Comparison of electron micrographssuggests that outer epithelium which secretes periostracum andmineral shell, is separated from inner epithelium by a bandof "lobate" cells, of variable width, exuding an impersistentmucopolysaccharide film or pellicle. The lobate cells alwaysoccupy the same relative position on the inner surface of theouter mantle lobe; but the outer epithelium is commonly connectedwith the inner surface of the periostracum by papillae and protoplasmicstrands which persist during mineral deposition and ensure thatboth shell and attached mantle remain in situ relative to theoutwardly expanding inner surface of the outer mantle lobe.In the prototypic brachiopod, the lobate cells are likely atfirst to have occupied the hinge of the mantel fold but laterto have been displaced into their present position by the rigidoutward growing edge of the mineral shell.     

A new genus and species of monostiliferous hoplonemertean (Nemertea: Enopla: Monostilifera) from Japan

A new genus and species of monostiliferous hoplonemertean, Diopsonemertes acanthocephala gen. et sp. nov., is described from Otsuchi Bay, Japan. Significant anatomical features of the new form include a body wall longitudinal musculature anteriorly divided into inner and outer layers by connective tissue, no pre-cerebral septum, the presence of a thin coat of diagonal muscle fibres between the body wall longitudinal and circular muscle layers in the foregut body region, cephalic retractor muscles derived only from the inner portion of the divided longitudinal muscles and a rhynchocoel more than half the body length.     

Stress gradients in the walls of large arteries

Elastic behavior of vascular wall, assuming the vessels to be ‘thick-walled’ and utilizing finite deformation theory, was investigated. It was found that canine carotid arterial wall is neither isotropic nor transversely isotropic. Previously, stress-strain relations were obtained for carotid arteries on the basis of membrane theory (Doyle and Dobrin, 1971). Since strain gradients across the wall are fairly steep, the applicability of such expressions, for pointwise evaluation of stress, required examination. The study indicated that these relationships between mean circumferential stress and mean extension ratio in the circumferential direction could be used to relate the specific circumferential stress value to the specific extension ratio at any designated point within the wall. From this analysis it was possible to evaluate circumferential and radial wall stresses. Both of these stresses are maximal at the inner surface of the intima. At this point the radial stress is equal to the transmural pressure and is compressive, while the circumferential stress is tensile and is 1·5 to 2 times the value of the mean stress, i.e. the product of transmural pressure and the ratio of internal radius-to-wall thickness. Both stresses are lowest at the outer edge of the adventitia. These stress distributions were considered with respect to the spacing of the elastic lamellae and the absence of discernible vasa vasora in the inner third of the wall.     

Morphology and wall ultrastructure of some Middle Devonian dispersed megaspores from northern Poland

Specimens of dispersed Middle Devonian megaspores have been isolated from core samples from the Miastko 1 borehole in Western Pomerania. Comprehensive investigations using light, scanning and transmission electron microscopy supplement previous information on morphology and gross structure and provide data on spore wall ultrastructure of four megaspore species. Corystisporites acutispinosus is azonate; the inner layer is laminate, and the lumen is lined by a thick, laterally continuous lamina. The outer layer consists of small, tangentially aligned tabular elements that become wider, more extensive and irregularly arranged toward the outside. Coronispora variabilis is a coronate megaspore; the inner layer appears homogenous and is probably lamellate. The outer layer consists of elongate, cylindrical, branching elements that are overlaid within the proximal part of the body by a lamellate, compact, almost homogenous layer. Grandispora ciliata is pseudosaccate. The inner body is laminate with laminae thickening and becoming less continuous and less tightly packed toward the outside. The outermost region of the inner body and the innermost region of the outer envelope consist of tabular and cylindrical elongate units. The bulk of the outer wall is almost homogenous, and near the surface it is granular. Pomeranisporites subtriangularis is pseudozonate. The inner layer appears homogenous except for the presence of a single innermost lamina. The inner part of the outer layer may represent small tabular and cylindrical elements, and the outer part comprises folded laminae. The megaspores studied share numerous features of morphology and wall ultrastructure with the lycopsids, putative lycopsids, and some enigmatic Devonian plants.     

The neuroactive substances and associated muscle system in Rhipidocotyle campanula (Digenea,Bucephalidae) from the intestine of the pike Esox lucius

We report about the muscular system and the serotonergic and FMRFamidergic components of the nervous system of the Bucephalidae trematode, Rhipidocotyle campanula, an intestinal parasite of the pike. We use immunocytochemical methods and confocal scanning laser microscopy (CLSM). The musculature is identified by histochemical staining with fluorescently labeled phalloidin. The body wall musculature of R. campanula contains three layers of muscle fibres – the outer thin circular, intermediate longitudinal and inner diagonal muscle fibres running in two opposite directions. The digestive system of R. campanula possess of a well-developed musculature: radial, longitudinal and circular muscle elements are detected in the pharynx, circular and longitudinal muscle filaments seen in the oesophagus, and longitudinal and the circular muscle fibres were found in the intestinal wall. Specific staining indicating the presence of actin muscle filaments occurs in the cirrus sac localized in the posterior body region. The frontal region of anterior attachment organ, the rhynchus, in R. campanula is represented by radial muscle fibres. The posterior part of the rhynchus comprise of radial muscles forming the organ's wall, and several strong longitudinal muscle bundles. Serotonergic and FMRFamidergic structures are detected in the central and peripheral compartments of the nervous system of R. campanula, that is, in the paired brain ganglia, the brain commissure, the longitudinal nerve cords, and connective nerve commissures. The innervations of the rhynchus, pharynx, oesophagus and distal regions of the reproductive system by the serotonergic and FMRFamidergic nervous elements are revealed. We compare our findings obtained on R. campanula with related data for other trematodes.     

Morphological studies on the mantle of the fresh-water mussel Amblema (UNIONIDAE): Scanning electron microscopy

The scanning electron microscope has been used to describe the surface morphology of the mantle in mantle-shell preparations from the fresh-water mussel Amblema. In some regions (adductor muscle insertions), the mantle is firmly attached to the shell. In other areas (along the main course of the mantle), transient adhesions between the outer mantle epithelial cells and the nacre appear to temporally further compartmentalize the extrapallial fluid possibly as a prerequisite for the initial crystallization phenomenon. At the mantle edge, as well as at the isthmus, the periostracum was seen to extrude from the periostracal groove. At the siphonal edge, peculiar fingerlike processes were identified; these may represent primitive photoreceptors. The epithelial cells of the outer mantle epithelium are all microvillated whereas those of the inner mantle epithelium are both microvillated and ciliated. Specific differences in surface morphology are described for various regions of the outer mantle epithelium. These may be related to precise regionalized functional differences of this tissue. Several functions of the mantle, in addition to shell formation, and based on its various morphologies, are also discussed.