Position of the labrum in agnostid trilobites

Specimens of Peronopsis and Ptychagnostus from the House Range of western Utah provide new information on the location of the labrum (hypostoma) of agnostid trilobites. A labral position as that described here, beneath the second glabellar lobe, has two implications: (1) anomalous cephalic muscle scars described by previous workers may be interpreted as attachment areas for labral muscles; and (2) the mouth probably migrated posteriorly in agnostids. Posterior displacement of the mouth was probably one of several feeding modifications associated with a pelagic mode of life.     

The insect upper lip (labrum) is a nonsegmental appendage-like structure

SUMMARY The insect upper lip—the labrum—is a lobe-like structure anterior to the mouth opening. Whether the labrum represents a fused pair of segmental appendages or evolved independently is heavily debated. Here, we identify additional similarities of the regulatory gene network active in labrum and trunk appendages. However, we do not find a labral parasegment boundary and we show that labral Tc-Dll expression is independent of Tc-wg and Tc-hh signals. In contrast, Tc-Dll expression in all trunk appendages does require these signals. Finally, we identify crucial differences between the location of the labrum and trunk appendages: the labrum develops in median rather than lateral tissues and is part of an anterior nonsegmental tissue marked by and dependent on Tc-six3 activity. To reconcile these seeming contradictory results, we propose that the genetic network evolved in either labrum or trunk appendages and became redeployed at a novel location to form the other structure.     

Homeotic evidence for the appendicular origin of the labrum in Tribolium castaneum

The ontogeny of the insect labrum, or upper lip, has been debated for nearly a century. Recent molecular data suggest a segmental appendage origin of this structure. Here we report the first arthropod mutation associated with a homeotic transformation of the labrum. Antennagalea-5 (Ag(5)) transforms both antennal and labral structures to resemble those of gnathal appendages in Tribolium castaneum. This labral transformation suggests that the labrum is a fused structure composed of two pairs of appendage endites, and is serially homologous to the gnathal appendages.     

Effect of acetabular labral tears,repair and resection on hip cartilage strain: A 7 T MR study

BackgroundTears of the acetabular labrum are frequently present in patients with groin pain. While it is clear that the labrum contributes to the surface area articulating with the femoral head, it is not clear whether labral repair yields different load distribution in the hip compared to labral resection.PurposeDetermine whether labral repair reduces cartilage strain more effectively than labral resection.MethodsSix human cadaveric hips (mean age 37 years) were loaded in a simulated single-leg stance within the bore of a 7 T MR scanner. After cartilage had reached a steady-state thickness distribution, a scan of the cartilage was acquired with a voxel size of 0.1×0.1×0.3 mm. This method was repeated for each of six specimens when the labrum was intact, after a surgically simulated labral tear, after an arthroscopic labral repair and after labral resection. Cartilage thickness and strain in an anterosuperior region of interest were measured from the MR scans. A paired t-test was used to compare mean and maximum cartilage strain when the labrum was intact vs. torn, torn vs. repaired and repaired vs. resected. Three-dimensional patterns of cartilage strain distribution were qualitatively compared for the different labral conditions.ResultsFor the number of specimens tested we found no change in mean and maximum cartilage strain, and little obvious change in the pattern of cartilage strain distribution after a simulated labral tear. Labral repair caused a 2% decrease in mean cartilage strain compared to a torn labrum (p=0.014). Labral resection caused a 4% and 6% increase in mean and maximum cartilage strain, respectively, compared to labral repair (p=0.02), and the cartilage strain distribution was elevated throughout the region of interest.ConclusionBased on our ex vivo findings of increased cartilage strain after labral resection when compared to labral repair, we have demonstrated the associated consequences to the mechanical environment of the cartilage following surgical treatment of the labrum.     

Ultrastructural investigations on the labral glands of Daphnia obtusa (Crustacea,Cladocera)

The labral glands of Daphnia consist of three distinct functional units on each side: (1) several cells at the base of the head, (2) two large cells at the base of the labrum and one large cell (cell A) in the median part of the labrum and (3) one large cell (cell B) in the median part of the labrum. These gland cells do not form a syncytium, contrary to reports by previous investigators. With the exception of cell B, they have a well-developed rough endoplasmic reticulum and many active Golgi complexes. The Golgi activity changes during the molt cycle. The Golgi activity of the cells of the head base is different from that of the large cells of the labrum. Since clear exocytotic phenomena were not observed, the secretion can be assumed to flow into the hemolymph after accumulation in the enlarged intercellular spaces. Cell B has a distinctive cytoplasmic ultrastructure the function of which is not yet understood. The four large cells of the labrum are in contact with a duct cell (or several duct cells) characterized by a deep infolding of the plasma membrane. This delimits a narrow lumen, which contains no secretion. No passage of substance is visible from the gland cells to the duct cell(s).     

Role of the acetabular labrum in load support across the hip joint

The relatively high incidence of labral tears among patients presenting with hip pain suggests that the acetabular labrum is often subjected to injurious loading in vivo. However, it is unclear whether the labrum participates in load transfer across the joint during activities of daily living. This study examined the role of the acetabular labrum in load transfer for hips with normal acetabular geometry and acetabular dysplasia using subject-specific finite element analysis. Models were generated from volumetric CT data and analyzed with and without the labrum during activities of daily living. The labrum in the dysplastic model supported 4-11% of the total load transferred across the joint, while the labrum in the normal model supported only 1-2% of the total load. Despite the increased load transferred to the acetabular cartilage in simulations without the labrum, there were minimal differences in cartilage contact stresses. This was because the load supported by the cartilage correlated with the cartilage contact area. A higher percentage of load was transferred to the labrum in the dysplastic model because the femoral head achieved equilibrium near the lateral edge of the acetabulum. The results of this study suggest that the labrum plays a larger role in load transfer and joint stability in hips with acetabular dysplasia than in hips with normal acetabular geometry.     

Formation of the arthropod labrum by fusion of paired and rotated limb-bud-like primordia

The nature and origin of the arthropod labrum is a matter much under dispute. We show here that in Tribolium castaneum (Herbst, 1797) the labrum develops from two individual primordia, termed labral buds. Expression of the genes decapentaplegic (dpp) and wingless (wg) in these buds is identical to the buds of the metameric appendages (e.g. thoracic legs), except that the patterns are reversed. We propose that this reversal is the result of the rotation of the labral buds through an angle of approximately 180°. We also for the first time study dpp and wg expression in the fully developed labrum of older embryonic stages. Here, gene expression patterns show that the labrum proper is formed by fusion of the labral buds along their dorsal sides, while their ventral sides are facing outward forming the lateral sides of the fused labrum. Furthermore, we show that there are very similar patterns in another arthropod species, the spider Cupiennius salei (Keyserling, 1877), although in this species the labrum develops as a single structure and not from two separate primordia. However, in C. salei the expression of engrailed is also reversed in addition to the reversal of dpp and wg expression: engrailed is expressed in the anterior half of the labrum, and not in the posterior half like in the remaining appendages. Our results suggest that the arthropod labrum is derived evolutionarily from paired limb-bud-like primordia by rotation and fusion, and that this process is recapitulated ontogenetically to a different extent in different arthropod species.     

Hip chondrolabral mechanics during activities of daily living: Role of the labrum and interstitial fluid pressurization

Osteoarthritis of the hip can result from mechanical factors, which can be studied using finite element (FE) analysis. FE studies of the hip often assume there is no significant loss of fluid pressurization in the articular cartilage during simulated activities and approximate the material as incompressible and elastic. This study examined the conditions under which interstitial fluid load support remains sustained during physiological motions, as well as the role of the labrum in maintaining fluid load support and the effect of its presence on the solid phase of the surrounding cartilage. We found that dynamic motions of gait and squatting maintained consistent fluid load support between cycles, while static single-leg stance experienced slight fluid depressurization with significant reduction of solid phase stress and strain. Presence of the labrum did not significantly influence fluid load support within the articular cartilage, but prevented deformation at the cartilage edge, leading to lower stress and strain conditions in the cartilage. A morphologically accurate representation of collagen fibril orientation through the thickness of the articular cartilage was not necessary to predict fluid load support. However, comparison with simplified fibril reinforcement underscored the physiological importance. The results of this study demonstrate that an elastic incompressible material approximation is reasonable for modeling a limited number of cyclic motions of gait and squatting without significant loss of accuracy, but is not appropriate for static motions or numerous repeated motions. Additionally, effects seen from removal of the labrum motivate evaluation of labral reattachment strategies in the context of labral repair.     

Shoulder labral pathomechanics with rotator cuff tears

Rotator cuff tears (RCTs), the most common injury of the shoulder, are often accompanied by tears in the superior glenoid labrum. We evaluated whether superior humeral head (HH) motion secondary to RCTs and loading of the long head of the biceps tendon (LHBT) are implicated in the development of this associated superior labral pathology. Additionally, we determined the efficacy of a finite element model (FEM) for predicting the mechanics of the labrum. The HH was oriented at 30° of glenohumeral abduction and neutral rotation with 50 N compressive force. Loads of 0 N or 22 N were applied to the LHBT. The HH was translated superiorly by 5 mm to simulate superior instability caused by RCTs. Superior displacement of the labrum was affected by translation of the HH (P<0.0001), position along the labrum (P<0.0001), and interaction between the location on the labrum and LHBT tension (P<0.05). The displacements predicted by the FEM were compared with mechanical tests from 6 cadaveric specimens and all were within 1 SD of the mean. A hyperelastic constitutive law for the labrum was a better predictor of labral behavior than the elastic law and insensitive to ±1 SD variations in material properties. Peak strains were observed at the glenoid–labrum interface below the LHBT attachment consistent with the common location of labral pathology. These results suggest that pathomechanics of the shoulder secondary to RCTs (e.g., superior HH translation) and LHBT loading play significant roles in the pathologic changes seen in the superior labrum.     

Embryonic development of the sensory innervation of the clypeo-labral complex: further support for serially homologous appendages in the locust

The clypeo-labrum, or upper lip, of insects is intimately involved in feeding behavior and is accordingly endowed with a rich sensory apparatus. In the present study we map the temporal appearance of all major clusters of sensory cells on this structure in the locust during the first half of embryogenesis. The identities of these sensory cell clusters were defined according to the origin of the branching point of their axons from the labral sensory nerve as seen at mid-embryogenesis. The first sensory cells to differentiate from the labral epithelium do so at stereotypic sites beginning at around 32% of embryogenesis. Bilaterally symmetrical clusters of differentiated neurons rapidly appear and pioneering of the labral sensory nerve on each side is performed by a specific cell from each cluster. This cell directs its axon anteriorly towards a bilaterally symmetrical pair of cells, the frontal commissure pioneers, on either side of the developing frontal ganglion. The final trajectory of the sensory nerve within the labrum closely matches the pattern of Repo-expressing glial cells. The majority of the sensory cell clusters differentiate during embryogenesis, but the number of sensory cells in some clusters are modified significantly during postembryonic development. Comparing the innervation pattern of the clypeo-labrum with that of other mouthparts and the leg at mid-embryogenesis, we find a striking similarity in organization which we interpret as support for the homologous appendage hypothesis.     

Embryonic origin of the imaginal discs of the head of Drosophila melanogaster

The embryonic development of the primordia of the Drosophila head was studied by using an enhancer trap line expressed in these structures from embryonic stage 13 onward. Particular attention was given to the question of how the adult head primordia relate to the larval head segments. The clypeo-labral bud to the stage 13 embryo is located at a lateral position in the labrum adjacent to the labral sensory complex (epiphysis). Both clypeo-labral bud and sensory complex are located anterior to the engrailed-expression domain of the labrum. Throughout late embryogenesis and the larval period, the clypeo-labral bud forms integral part of the epithelium lining the roof of the atrium. The labial disc originates from the lateral labial segment adjacent to the labial sensory complex (hypophysis). It partially overlaps with the labial en-domain. After head involution, the labial disc forms a small pocket in the ventro-lateral wall of the atrium. The eye-antenna disc develops from a relatively large territory occupying the dorso-posterior part of the procephalic lobe, as well as parts of the dorsal gnathal segments. Cells in this territory are greatly reduced in number by cell death during stages 12–14. After head involution, the presumptive eye-antenna disc occupies a position in the lateral-posterior part of the dorsal pouch. Evagination of this tissue occurs during the first hours after hatching. In the embryo, no en-expression is present in the presumptive eye-antenna disc. en-expression starts in three separate regions in the third instar larva.     

Morphological and molecular data argue for the labrum being non-apical, articulated, and the appendage of the intercalary segment in the locust

Our analysis of head segmentation in the locust embryo reveals that the labrum is not apical as often interpreted but constitutes the topologically fused appendicular pair of appendages of the third head metamere. Using molecular, immunocytochemical and retrograde axonal staining methods we show that this metamere, the intercalary segment, is innervated by the third brain neuromere-the tritocerebrum. Evidence for the appendicular nature of the labrum is firstly, the presence of an engrailed stripe within its posterior epithelium as is typical of all appendages in the early embryo. Secondly, the labrum is innervated by a segmental nerve originating from the third brain neuromere (the tritocerebrum). Immunocytochemical staining with Lazarillo and horseradish peroxidase antibodies reveal that sensory neurons on the labrum contribute to the segmental (tritocerebral) nerve via the labral nerve in the same way as for the appendages immediately anterior (antenna) and posterior (mandible) on the head. All but one of the adult and embryonic motoneurons innervating the muscles of the labrum have their cell bodies and dendrites located completely within the tritocerebral neuromere and putatively derive from engrailed expressing tritocerebral neuroblasts. Molecular evidence (repo) suggests the labrum is not only appendicular but also articulated, comprising two jointed elements homologous to the coxa and trochanter of the leg.     

Two microvillar organs, new to Crustacea, in the Mystacocarida

The mystacocarid crustacean Derocheilocaris typica has two microvillar organs, one new, the other previously unappreciated in crustacean literature. The first is situated on the head-shield and consists of three pairs of cells: one with microvilli and a ballooned nucleus; one smaller and without special features; the third large and investing the other two and extending down to the foregut. We call this new organ the "cephalic microvillar organ" and discuss the value of the concept "dorsal organ", to which it might have been included. The second organ consists of about 21 cells that cover the proximal part of the dorsal surface of the labrum. The cells are alike, being characterized by an apical field of microvilli and a large residual body. This organ is here called the "labral microvillar organ". Both organs are neither sensory nor secretory and do not qualify for membership in any of the other recognized organ systems. We are unable to deduce their Dero-cheilocaris functions.     

The mechanics of focal chondral defects in the hip

There is a mean incidence of osteoarthritis (OA) of the hip in 8% of the overall population. In the presence of focal chondral defects, defined as localized damage to the articular cartilage, there is an increased risk of symptomatic progression toward OA. This relationship between chondral defects and subsequent development of OA has led to substantial efforts to develop effective procedures for surgical cartilage repair. This study examined the effects of chondral defects and labral delamination on cartilage mechanics in the dysplastic hip during the gait cycle using subject-specific finite element analysis. Models were generated from volumetric CT data and analyzed with simulated chondral defects at the chondrolabral junction on the posterior acetabulum during five distinct points in the gait cycle. Focal chondral defects increased maximum shear stress on the osteochondral surface of the acetabular cartilage, when compared to the intact case. This effect was amplified with labral delamination. Additionally, chondral defects increased the first principal Lagrange strain on the articular surface of the acetabular cartilage and labrum. Labral delamination relieved some of this tensile strain. As defect size was increased, contact stress increased in the medial zone of the acetabulum, while it decreased anteriorly. The results suggest that in the presence of chondral defects and labral delamination the cartilage experiences elevated tensile strains and shear and contact stress, which could lead to further damage of the cartilage, and subsequent arthritic progression. The framework presented here will serve as the procedure for future finite element studies on cartilage mechanics in hips with varying disease states with simulated chondral defects and labral tears.     

Anatomy of membranous mouthpart cuticles and their roles in feeding in black flies (Diptera: Simuliidae)

The nature and extent of the cuticular membranes of the mouthparts of females of Simuiium venustum are described. These membranes are arranged into four groups: (1) cuticular membranes of the labium/hypopharynx; (2) membranes of the apical labrum; (3) those of basal portions of the mandibles and laciniae; and (4) the labral flaps within the syntrophial food canal. From their external and internal structure it is concluded that membranes of the first three groups go through inflation/deflation cycles directly related to the food-pumping rhythm. It is further concluded that these membranes line the inner and outer syntrophial and proboscis surfaces, acting as pressure seals both to prevent blood loss from the wound and air entry into the food canal during feeding. The labral flaps within the food canal appear to act as one-way valves, keeping blood from leaking out between the mouthparts during the pumping downstroke but allowing pooled blood to enter during the upstroke.     

Molecular phylogenetics and the evolution of labral spines among eastern Pacific ocenebrine gastropods

Labral spines are sharp projections of the apertural lip found in some marine gastropods that are used to penetrate hard-shelled prey. The majority of gastropod genera that contain labral spine-bearing species are found in the subfamily Ocenebrinae (Gastropoda: Muricidae). To reconstruct the evolutionary history of labral spine-bearing and labral spine-lacking gastropods in the eastern Pacific (EP) Ocean, partial sequences of two mitochondrial genes (cytochrome oxidase I and 12S rRNA) were obtained from representative taxa. Despite high nucleotide bias, a variety of phylogenetic reconstruction methods produced the same tree topology. The traditional taxonomic view that all "Nucella-like" spine-bearing taxa in the EP belong to a monophyletic "Acanthina" is rejected due to nonmonophyly of this group. The more recently recognized "Acanthinucella" is also not monophyletic, and we therefore propose the new genus Mexacanthina for two Mexican species formerly assigned to Acanthinucella. The genus Ocinebrina, which first appears in the middle Eocene, is not a stem EP ocenebrine lineage and may also not be a monophyletic clade. Tracing the evolutionary history of labral spines among extant lineages indicates that the absence of a labral spine is ancestral for all EP ocenebrines. Ancestral conditions could not be resolved unambiguously for all nodes of the phylogeny based on extant taxa. However, by jointly considering both molecular phylogenetic relationships and the phylogenetic affinities of several extinct taxa, all remaining character state transformation can be inferred unambiguously. Based on this analysis, a labral spine likely evolved independently in at least four lineages of EP ocenebrines. Although homoplasy appears to characterize labral spine evolution among ocenebrine gastropods, the structural position of a labral spine was evolutionarily altered in one lineage, indicating that different types of labral spines do not necessarily reflect convergent evolution.     

Composite,haustellate mouthparts in netwinged beetle and firefly larvae (Coleoptera,Cantharoidea: Lycidae,Lampyridae)

During lycid embryogenesis, labral and mandibular anlagen develop separately. Each anterolateral corner of the labrum elongates into a stylet, while the whole of each mandible involutes into a flute that is open to the lateral aspects. The stylets and flutes interlock in such a way as to leave canals, fashioning left and right haustellate apparatuses. During lampyrid embryogenesis, only one pair of anlagen forms instead of two. The pair matures into sharp, arcuate appendages that appear to be typical mandibles, but nevertheless a canal forms internally. The canal is not formed by coadaptation of two appendages; instead, cells within the pair create the canal autonomously. Indications are that the canal is homologous between these two families. It is suggested that in lampyrids, labral and mandibular cell fields develop together in the same anlage but behave according to their own respective programs, forming the canal with morphogenetic movements that are comparable to those of lycids. © 1994 Wiley-Liss, Inc.     

Endonura Cassagnau in Iran,with a key to species of the genus (Collembola,Neanuridae, Neanurinae)

Three new species of Endonura are described from Iran. Endonura dichaeta sp. n. can be recognized by an ogival labrum, head without chaetae O and E, chaeta D connected with tubercle Cl, tubercle Dl with five chaetae on head, absence of tubercles Di on thorax I and tubercle (Di+Di) of thorax V with 2+2 chaetae. Endonura ceratolabralis sp. n. is characterized by large body size, reduction of labral chaetotaxy, ogival labrum, head without chaeta O and fusion of tubercles Di and De on first thoracic segment. Endonura persica sp. n. is distinguished from its congeners by a nonogival labrum, absence of chaeta O, tubercles Dl and (L+So) with five and eight chaetae respectively and claw with inner tooth. The key to all species of the genus is given.     

Development of the mouthparts in embryos of Haplothrips verbasci (Osborn) (Insecta,Thysanoptera, Phlaeothripidae)

The asymmetric “punch and suck” mouthparts of larval Haplothrips verbasci develop from paired appendages in the late, post-anatrepsis embryo similar to those of other insects. Later, the labrum flexes ventrally over the stomodaeum, the right mandibular appendage degenerates, the maxillary appendages divide into inner (lacinial) and outer (stipital) lobes, and the hypopharynx arises from the venters of the mandibular and maxillary segments. All cephalic segments consolidate anteriorly prior to katatrepsis, their appendages flex ventrally, and the labial appendages fuse medially to form the labium and the primordia of the salivary glands and valve. The left mandible and the lacinial lobes of the maxillae invaginate into the head during and after katatrepsis to form the mandibular and maxillary stylet-secreting organs and these later deposit the cuticle of their respective stylets. Cuticle of the mandibular lever is deposited by labral cells at the apex of the mandibular sheath during and after hatching. That of each maxillary lever is secreted simultaneously into the lumen of a ventrally-directed diverticulum developing from stipital cells at the apex of each maxillary sheath. Shortly after katatrepsis, the maxillary and labial palpi originate respectively from cells in the outer wall of each stipital lobe and at the apex of the labium. Muscles of the mouthparts arise after katatrepsis from cephalic mesoderm and are fully-differentiated before cuticle of the mandibular and maxillary levers has been deposited. Gnathal morphogenesis in embryos of H. verbasci resembles that occurring in bug embryos and provides additional evidence that Thysanoptera and Hemiptera evolved from a common psocopteroid stem species having small, paired, biting and chewing mandibles and well developed lacinial stylets.