PubMed Health⌕ Search

Biomedical subjects

E L Brainerd

Publications and source records attributed to E L Brainerd.

11 recordsLinked to original sources

Patterns of genome size evolution in tetraodontiform fishes.

We used flow cytometry to measure genome size in 15 species from seven families and subfamilies of tetraodontiform fishes. Previous studies have found that smooth pufferfishes (Tetraodontidae) have the smallest genome of any vertebrate measured to date (0.7-1.0 picograms diploid). We found that spiny pufferfishes (Diodontidae, sister group to the smooth puffers) possess a genome that is about two times larger (1.6-1.8 pg). Mola mola, a member of the sister group to Diodontidae and Tetraodontidae, also has a relatively large genome (1.7 pg). Parsimony analysis of this pattern indicates that the plesiomorphic condition for Molidae (Diodontidae, Tetraodontidae) is a genome size of 1.6-1.8 pg, and that tiny genome size is a derived character unique to smooth puffers. However, an alternative explanation is that the ancestor of Tetraodontidae acquired a heritable tendency toward decreasing genome size, such as a new or modified deletion mechanism, and genome size in all of the tetraodontid lineages has been decreasing in parallel since the split from Diodontidae. Small genome size (1.1-1.3 pg) also appears to have evolved independently in some members of Balistoidea (triggerfishes and filefishes) within Tetraodontiformes.

Animals↗

Biomechanics of the movable pretarsal adhesive organ in ants and bees.

Hymenoptera attach to smooth surfaces with a flexible pad, the arolium, between the claws. Here we investigate its movement in Asian weaver ants (Oecophylla smaragdina) and honeybees (Apis mellifera). When ants run upside down on a smooth surface, the arolium is unfolded and folded back with each step. Its extension is strictly coupled with the retraction of the claws. Experimental pull on the claw-flexor tendon revealed that the claw-flexor muscle not only retracts the claws, but also moves the arolium. The elicited arolium movement comprises (i) about a 90 degrees rotation (extension) mediated by the interaction of the two rigid pretarsal sclerites arcus and manubrium and (ii) a lateral expansion and increase in volume. In severed legs of O. smaragdina ants, an increase in hemolymph pressure of 15 kPa was sufficient to inflate the arolium to its full size. Apart from being actively extended, an arolium in contact also can unfold passively when the leg is subject to a pull toward the body. We propose a combined mechanical-hydraulic model for arolium movement: (i) the arolium is engaged by the action of the unguitractor, which mechanically extends the arolium; (ii) compression of the arolium gland reservoir pumps liquid into the arolium; (iii) arolia partly in contact with the surface are unfolded passively when the legs are pulled toward the body; and (iv) the arolium deflates and moves back to its default position by elastic recoil of the cuticle.

Animals↗

Buccal oscillation and lung ventilation in a semi-aquatic turtle, Platysternon megacephalum.

Movements of the hyobranchial apparatus in reptiles and amphibians contribute to many behaviors including feeding, lung ventilation, buccopharyngeal respiration, thermoregulation, olfaction, defense and display. In a semi-aquatic turtle, Platysternon megacephalum, x-ray video and airflow measurements from blowhole pneumotachography show no evidence that above water hyobranchial movements contribute to lung inflation, as in the buccal or gular pump of amphibians and some lizards. Instead, hyobranchial movements produce symmetrical oscillations of air into and out of the buccal cavity. The mean tidal volume of these buccal oscillations is 7.8 times smaller than the mean tidal volume of lung ventilation (combined mean for four individuals). Airflow associated with buccal oscillation occurs in the sequence of inhalation followed by exhalation, distinguishing it from lung ventilation which occurs as exhalation followed by inhalation. No fixed temporal relationship between buccal oscillation and lung ventilation was observed. Periods of ventilation often occur without buccal oscillation and buccal oscillation sometimes occurs without lung ventilation. When the two behaviors occur together, the onset of lung ventilation often interrupts buccal oscillation. The initiation of lung ventilation was found to occur in all phases of the buccal oscillation cycle, suggesting that the neural control mechanisms of the two behaviors are not coupled. The pattern of occurrence of both buccal oscillation and lung ventilation was found to vary over time with no obvious effect of activity levels.

Journal Article↗

Twisting and bending: the functional role of salamander lateral hypaxial musculature during locomotion.

The function of the lateral hypaxial muscles during locomotion in tetrapods is controversial. Currently, there are two hypotheses of lateral hypaxial muscle function. The first, supported by electromyographic (EMG) data from a lizard (Iguana iguana) and a salamander (Dicamptodon ensatus), suggests that hypaxial muscles function to bend the body during swimming and to resist long-axis torsion during walking. The second, supported by EMG data from lizards during relatively high-speed locomotion, suggests that these muscles function primarily to bend the body during locomotion, not to resist torsional forces. To determine whether the results from D. ensatus hold for another salamander, we recorded lateral hypaxial muscle EMGs synchronized with body and limb kinematics in the tiger salamander Ambystoma tigrinum. In agreement with results from aquatic locomotion in D. ensatus, all four layers of lateral hypaxial musculature were found to show synchronous EMG activity during swimming in A. tigrinum. Our findings for terrestrial locomotion also agree with previous results from D. ensatus and support the torsion resistance hypothesis for terrestrial locomotion. We observed asynchronous EMG bursts of relatively high intensity in the lateral and medial pairs of hypaxial muscles during walking in tiger salamanders (we call these 'alpha-bursts'). We infer from this pattern that the more lateral two layers of oblique hypaxial musculature, Mm. obliquus externus superficialis (OES) and obliquus externus profundus (OEP), are active on the side towards which the trunk is bending, while the more medial two layers, Mm. obliquus internus (OI) and transversus abdominis (TA), are active on the opposite side. This result is consistent with the hypothesis proposed for D. ensatus that the OES and OEP generate torsional moments to counteract ground reaction forces generated by forelimb support, while the OI and TA generate torsional moments to counteract ground reaction forces from hindlimb support. However, unlike the EMG pattern reported for D. ensatus, a second, lower-intensity burst of EMG activity ('beta-burst') was sometimes recorded from the lateral hypaxial muscles in A. tigrinum. As seen in other muscle systems, these beta-bursts of hypaxial muscle coactivation may function to provide fine motor control during locomotion. The presence of asynchronous, relatively high-intensity alpha-bursts indicates that the lateral hypaxial muscles generate torsional moments during terrestrial locomotion, but it is possible that the balance of forces from both alpha- and beta-bursts may allow the lateral hypaxial muscles to contribute to lateral bending of the body as well.

Ambystoma↗

Mechanics of lung ventilation in a post-metamorphic salamander, Ambystoma Tigrinum.

The mechanics of lung ventilation in frogs and aquatic salamanders has been well characterized, whereas lung ventilation in terrestrial-phase (post-metamorphic) salamanders has received little attention. We used electromyography (EMG), X-ray videography, standard videography and buccal and body cavity pressure measurements to characterize the ventilation mechanics of adult (post-metamorphic) tiger salamanders (Ambystoma tigrinum). Three results emerged: (i) under terrestrial conditions or when floating at the surface of the water, adult A. tigrinum breathed through their nares using a two-stroke buccal pump; (ii) in addition to this narial two-stroke pump, adult tiger salamanders also gulped air in through their mouths using a modified two-stroke buccal pump when in an aquatic environment; and (iii) exhalation in adult tiger salamanders is active during aquatic gulping breaths, whereas exhalation appears to be passive during terrestrial breathing at rest. Active exhalation in aquatic breaths is indicated by an increase in body cavity pressure during exhalation and associated EMG activity in the lateral hypaxial musculature, particularly the M. transversus abdominis. In terrestrial breathing, no EMG activity in the lateral hypaxial muscles is generally present, and body cavity pressure decreases during exhalation. In aquatic breaths, tidal volume is larger than in terrestrial breaths, and breathing frequency is much lower (approximately 1 breath 10 min(-)(1 )versus 4-6 breaths min(-)(1)). The use of hypaxial muscles to power active exhalation in the aquatic environment may result from the need for more complete exhalation and larger tidal volumes when breathing infrequently. This hypothesis is supported by previous findings that terrestrial frogs ventilate their lungs with small tidal volumes and exhale passively, whereas aquatic frogs and salamanders use large tidal volumes and and exhale actively.

Ambystoma↗

Contribution of gular pumping to lung ventilation in monitor lizards.

A controversial hypothesis has proposed that lizards are subject to a speed-dependent axial constraint that prevents effective lung ventilation during moderate- and high-speed locomotion. This hypothesis has been challenged by results demonstrating that monitor lizards (genus Varanus) experience no axial constraint. Evidence presented here shows that, during locomotion, varanids use a positive pressure gular pump to assist lung ventilation. Disabling the gular pump reveals that the axial constraint is present in varanids but it is masked by gular pumping under normal conditions. These findings support the prediction that the axial constraint may be found in other tetrapods that breathe by costal aspiration and locomote with a lateral undulatory gait.

Animals↗

Morphological variation of hypaxial musculature in salamanders (Lissamphibia: caudata).

Despite the acknowledged importance of the locomotory and respiratory functions associated with hypaxial musculature in salamanders, variation in gross morphology of this musculature has not been documented or evaluated within a phylogenetic or ecological context. In this study, we characterize and quantify the morphological variation of lateral hypaxial muscles using phylogenetically and ecologically diverse salamander species from eight families: Ambystomatidae (Ambystoma tigrinum), Amphiumidae (Amphiuma tridactylum), Cryptobranchidae (Cryptobranchus alleganiensis), Dicamptodontidae (Dicamptodon sp.), Plethodontidae (Gyrinophilus porphyriticus), Proteidae (Necturus maculosus), Salamandridae (Pachytriton sp.), and Sirenidae (Siren lacertina). For the lateral hypaxial musculature, we document 1) the presence or absence of muscle layers, 2) the muscle fiber angles of layers at mid-trunk, and 3) the relative dorsoventral positions and cross-sectional areas of muscle layers. Combinations of two, three, or four layers are observed. However, all species retain at least two layers with opposing fiber angles. The number of layers and the presence or absence of layers vary within species (Necturus maculosus and Siren lacertina), within genera (e.g., Triturus), and within families. No phylogenetic pattern in the number of layers can be detected with a family-level phylogeny. Fiber angle variation of hypaxial muscles is considerable: fiber angles of the M. obliquus externus range from 20-80 degrees; M. obliquus internus, 14-34 degrees; M. transversus abdominis, 58-80 degrees (acute angles measured relative to the horizontal septum). Hypaxial musculature comprises 17-37% of total trunk cross-sectional area. Aquatic salamanders show relatively larger total cross-sectional hypaxial area than salamanders that are primarily terrestrial.

Anatomy, Cross-Sectional↗

Kinematics of aquatic and terrestrial prey capture in Terrapene carolina, with implications for the evolution of feeding in cryptodire turtles.

Studies of aquatic prey capture in vertebrates have demonstrated remarkable convergence in kinematics between diverse vertebrate taxa. When feeding in water, most vertebrates employ large-amplitude hyoid depression to expand the oral cavity and suck in water along with the prey. In contrast, vertebrates feeding on land exhibit little or no hyoid depression. In this study we compared the kinematics of terrestrial and aquatic prey capture within one species of turtle, Terrapene carolina, in order to determine whether an individual species can modulate the magnitude of hyoid depression between air and water. High-speed video (250 frames per second) showed that hyoid depression was over three times greater in aquatic than in terrestrial feedings, indicating that T. carolina is able to modulate hyoid depression magnitude depending on the medium in which feeding occurs. In addition, we observed medium-dependent modulation of hyoid depression in another turtle, Heosemys grandis, and large-amplitude hyoid depression during aquatic feeding in Kinosternon leucostomum, Platysternon megacephalum, and juvenile Chelydra serpentina. In all of these turtles, hyoid depression produced oral cavity expansion during aquatic feeding, but the earthworm prey were never sucked toward the predators. Prey were captured by neck extension (ram feeding), and we conclude that the function of hyoid depression during aquatic feeding in cryptodire turtles is to prevent the forward motion of the predator from pushing the prey away (compensatory suction). Aquatic feeding is probably the primitive condition for all extant turtles, and thus terrestrial feeding in T. carolina and other turtles is a secondarily derived characteristic. We conclude from this historical pattern that it is not appropriate to use extant turtles in attempts to reconstruct the terrestrial feeding mechanisms of primitive amniotes.

Animals↗

Mechanics of lung ventilation in a larval salamander Ambystoma tigrinum.

The larval stage of the tiger salamander Ambystoma tigrinum is entirely aquatic, but the larvae rely on their lungs for a large proportion of their oxygen uptake. X-ray video and pressure measurements from the buccal and body cavities demonstrate that the larvae inspire using a two-stroke buccal pump and exhale actively by contracting the hypaxial musculature to increase body pressure. Larvae begin a breath by expanding the buccal cavity to draw in air through the mouth, while simultaneously exhaling air from the lungs to mix with the fresh air in the buccal cavity. The mouth then closes, and the buccal cavity compresses to pump a portion of the mixture into the lungs. The remaining air in the buccal cavity is then released as bubbles from the mouth and gill slits. Ventilatory volumes estimated from X-ray video records indicate that approximately 80% of the air pumped into the lungs is fresh air and 20% is previously expired air. Exhalation in larval tiger salamanders is active, powered by contraction of all four layers of lateral hypaxial musculature. Electromyography indicates that the transverse abdominis (TA) muscle is active for the longest duration and shows the highest-amplitude activity, but the external oblique superficialis, the external oblique profundus and the internal oblique also show consistent, low-level activity. The finding that the TA muscle is active during exhalation in larval tiger salamanders contributes to a growing body of evidence that the use of the TA for exhalation is a primitive character for tetrapods.

Ambystoma↗