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Biomedical subjects

Adam P Summers

Publications and source records attributed to Adam P Summers.

13 recordsLinked to original sources

Mineralized cartilage in the skeleton of chondrichthyan fishes.

The cartilaginous endoskeleton of chondrichthyan fishes (sharks, rays, and chimaeras) exhibits complex arrangements and morphologies of calcified tissues that vary with age, species, feeding behavior, and location in the body. Understanding of the development, evolutionary history and function of these tissue types has been hampered by the lack of a unifying terminology. In order to facilitate reciprocal illumination between disparate fields with convergent interests, we present levels of organization in which crystal orientation/size delimits three calcification types (areolar, globular, and prismatic) that interact in two distinct skeletal types, vertebral and tessellated cartilage. The tessellated skeleton is composed of small blocks (tesserae) of calcified cartilage (both prismatic and globular) overlying a core of unmineralized cartilage, while vertebral cartilage usually contains all three types of calcification.

Animals↗

The retro-articular process, streptostyly and the caecilian jaw closing system.

Caecilians have two functionally separate sets of jaw closing muscles. The jaw adductor muscles are parallel fibered muscles positioned close to the jaw joint and their lever mechanics suggests they are well suited to rapidly closing the jaws. A second set of muscles, the hypaxial interhyoideus posterior (IHP), levers the jaws closed by pulling on the retro-articular process (RA) of the lower jaw. Models of the lower jaw point out that the angle and length of the RA has a profound effect on the closure force exerted by the IHP. The caecilian skull is streptostylic - the quadrate-squamosal apparatus (QSA) moves relative to the rest of the skull, a condition that seems at odds with a well-ossified cranium. Modeling the contribution of this streptostylic suspension of the lower jaw shows that rotational freedom of the QSA amplifies the force of the IHP by redirecting force applied along the low axis of the lower jaw. Measurements from several species and life stages of preserved caecilians reveal a large variation in predicted bite force (as a multiple of IHP force) with age and phylogeny.

Amphibians↗

Gumfooted lines in black widow cobwebs and the mechanical properties of spider capture silk.

Orb-weaving spiders produce webs using two types of silk that have radically different mechanical properties. The dragline silk used to construct the supporting frame and radii of the web is stiff and as strong as steel, while the capture spiral is much weaker but more than ten times as extensible. This remarkable divergence in mechanical properties has been attributed to the aqueous glue that coats the capture spiral, which is thought to decrease capture spiral stiffness and increase its extensibility. However, discerning the effect of the aqueous glue on fiber performance is complicated because dragline silk and the capture spiral are assembled from different proteins, which may also affect mechanical performance. Here, we use the sticky gumfooted lines of black widow cobwebs to test the effect of the addition of aqueous glue on the mechanical properties of dragline silk. We also surveyed orb-webs spun by a broad range of species for bundles of looped silk. Such bundles, termed windlasses, have been thought to increase capture spiral extensibility by "paying out" additional lengths of silk. Our results suggest that neither plasticization of silk by aqueous glue nor excess silk in windlasses can by themselves account for the remarkable extensibility of orb-weaver capture silk compared to other spider silks. This argues that the unique amino acid motifs of the flagelliform fibroins that constitute the core of the capture spiral play an essential role in capture silk's extreme extensibility.

Amino Acid Motifs↗

Olfactory morphology of carcharhinid and sphyrnid sharks: does the cephalofoil confer a sensory advantage?

Many hypotheses have been advanced to explain the adaptive significance of the sphyrnid cephalofoil, including potential advantages of spacing the olfactory organs at the distal tips of the broad surface. We employed comparative morphology to test whether the sphyrnid cephalofoil provides better stereo-olfaction, increases olfactory acuity, and samples a greater volume of the medium compared to the situation in carcharhiniform sharks. The broadly spaced nares provide sphyrnid species with a significantly greater separation between the olfactory rosettes, which could lead to an enhanced ability to resolve odor gradients. In addition, most sphyrnid species possess prenarial grooves that greatly increase the volume of water sampled by the nares and thus increase the probability of odorant encounter. However, despite a much greater head width, and a significantly greater number of olfactory lamellae, scalloped hammerhead sharks do not possess a greater amount of olfactory epithelial surface area than the carcharhiniform sandbar sharks. Therefore, sphyrnid sharks might not possess any greater olfactory acuity than carcharhinids. Despite this, there are clear olfactory advantages to the cephalofoil head morphology that could have led to its evolution, persistence, and diversification. persistence, and diversification.

Animals↗

Batoid wing skeletal structure: novel morphologies, mechanical implications, and phylogenetic patterns.

The skeleton of the "wings" of skates and rays consists of a series of radially oriented cartilaginous fin rays emanating from a modified pectoral girdle. Each fin ray consists of small, laterally oriented skeletal elements, radials, traditionally represented as simple cylindrical building blocks. High-resolution radiography reveals the pattern of calcification in batoid wing elements, and their organization within the fin ray, to be considerably more complex and phylogenetically variable than previously thought. Calcification patterns of radials varied between families, as well as within individual pectoral fins. Oscillatory swimmers show structural interconnections between fin rays in central areas of the wing. Morphological variation was strongly predictive of locomotor strategy, which we attribute to oscillatory swimmers needing different areas of the wing stiffened than do undulatory swimmers. Contributions of various forms of calcification to radial stiffness were calculated theoretically. Results indicate that radials completely covered by mineralized tissue ("crustal calcification") were stiffer than those that were calcified in chain-like patterns ("catenated calcification"). Mapping this functionally important variation onto a phylogeny reveals a more complicated pattern than the literature suggests for the evolution of locomotor mode. Therefore, further investigation into the phylogenetic distribution of swimming mode is warranted.

Animals↗

The sexually dimorphic cephalofoil of bonnethead sharks, Sphyrna tiburo.

Sexually dimorphic head shape is common in vertebrates from teleosts to mammals. Herein we document that cephalic sexual dimorphism is also found in the cartilaginous fishes (Chondrichthyes). Male bonnethead sharks develop a prominent bulge along the anterior margin of the cephalofoil at the onset of sexual maturity. This contrasts with the uniformly rounded anterior margin of adult females and juveniles and embryos of both sexes. The anterior cephalic bulge is produced by elongation of the rod-like rostral cartilages, and its appearance corresponds temporally with the elongation of the rod-like cartilages of the male intromittent organs (claspers). We propose that the rostral cartilage elongation is a byproduct of endocrinological changes at the onset of sexual maturity that stimulate growth of the clasper cartilages. The basal location of the chondrichthyan fishes within the vertebrate clade extends the earliest appearance of cephalic sexual dimorphism among the vertebrates.

Animals↗

Structure and function of the horn shark (Heterodontus francisci) cranium through ontogeny: development of a hard prey specialist.

The horn sharks (Heterodontidae: Chondrichthyes) represent one of four independent evolutions of durophagy in the cartilaginous fishes. We used high-resolution computed tomography (CT scanning) to visualize and quantify the mineralized tissue of an ontogenetic series of horn sharks. CT scanning of neonatal through adult California horn sharks (Heterodontus francisci) confirmed that this technique is effective for examining mineralized tissue in even small (<10 mm) specimens. The jaw joint is among the first areas to become mineralized and is the most heavily mineralized area in the cranium of a neonatal horn shark. The hyoid is also well mineralized, although the poorly mineralized molariform teeth indicate that the neonatal animal may be a suction feeder on softer prey. The symphysis of the jaws never mineralizes, in sharp contrast to the condition in the hard prey-crushing stingrays. Digitally reslicing the CT scans along the jaws allowed measurement of the second moment of area (Ina). Assuming that the jaws are made of the same material at all ages, Ina is an indicator of the flexural stiffness of the jaws. In all sizes of shark the lower jaws were stiffer than the upper and the stiffness increased in the area of the molariform teeth. The central region of the jaws, where the rami meet, support cuspidate grasping teeth and has the lowest Ina. The spotted eagle ray (Aetobatus narinari), a hard prey-crushing stingray, shows a different pattern of flexural stiffness, with the peak at the central part of the jaws where the prey is reduced between flattened tooth plates. Although the eagle ray jaws have a higher Ina than the horn shark, they are also far more heavily mineralized. When the relative amounts of mineralization are taken into account, horn sharks do better with what mineral they have than does the eagle ray. With a tight jaw joint and loose mandibular symphysis, as well as nearly opposite patterns of stiffness in the jaws, it is clear that two of the clades of hard prey specialists use very different methods for cracking the hard prey problem.

Animals↗

A novel fibrocartilaginous tendon from an elasmobranch fish ( Rhinoptera bonasus).

Tendons of the jaw adductor muscles of a hard prey crushing stingray exhibit similar adaptations to compressive and shear loads as those seen in mammalian tendons. Ventral intermandibular tendon from the cownose ray, Rhinoptera bonasus, has a prominent fibrocartilaginous pad that lies between a fibrous region of the tendon and the mineralized tissue of the jaw. Histologically the pad is similar to the fibrocartilaginous meniscus of mammals, and these tissues also share some biochemical traits. Proteoglycan (PG) content in the fibrocartilaginous pad is nearly four times higher than in the linearly arrayed tendinous tissue. The predominant PGs appear to be an aggrecan-like molecule and a decorin-like molecule. The decorin-like molecule is quite small when compared to mammalian decorin (20-80 kDa vs. 100-200 kDa). This study is the first to document adaptations to compressive/shear loading in tendon from a cartilaginous fish, and the similarities to the mammalian condition argue for the early evolution of this reactive ability of tendinous tissue.

Animals↗

Maneuvering in juvenile carcharhinid and sphyrnid sharks: the role of the hammerhead shark cephalofoil.

The peculiar head morphology of hammerhead sharks has spawned a variety of untested functional hypotheses. One of the most intuitively appealing ideas is that the anterior foil acts, as in canard-winged aircraft, to increase maneuverability. We tested this hypothesis by determining whether juveniles of two hammerhead species (Sphyrna tiburo and S. lewini) turn more sharply, more often, and with greater velocity than a juvenile carcharhinid shark (Carcharhinus plumbeus). Although the hammerheads were more maneuverable, further investigation revealed that they do not roll their body during turns, suggesting that the cephalofoil does not act as a steering wing. We also show that hammerhead sharks demonstrate greater lateral flexure in a turn than carcharhinids, and that this flexibility may be due to cross sectional shape rather than number of vertebrae.

Journal Article↗

The evolution of tendon--morphology and material properties.

Phylogenetically, tendinous tissue first appears in the invertebrate chordate Branchiostoma as myosepta. This two-dimensional array of collagen fibers is highly organized, with fibers running along two primary axes. In hagfish the first linear tendons appear and the myosepta have developed specialized regions with unidirectional fiber orientation-a linear tendon within the flat sheet of myoseptum. Tendons react to compressive load by first forming a fibrocartilaginous pad, and under severe stress, sesamoid bones. Evidence for this ability to react to load first arises in the cartilaginous fish, here documented in a tendon from the jaw of a hard-prey crushing stingray. Sesamoid bones are common in bony fish and also in tetrapods. Tendons will also calcify under tensile loads in some groups of birds, and this reaction to load is seen in no other vertebrates. We conclude that the evolutionary history of tendon gives us insight into the use of model systems for investigating tendon biology. Using mammal and fish models may be more appropriate than avian models because of the apparent evolution of a novel reaction to tensile loads in birds.

Animals↗