Dynamic changes in body form during swimming in the water snake Nerodia sipedon

Video records of swimming water snakes show that during moderate to rapid swimming, the rear half to two-thirds of the trunk is compressed laterally, approaching the body form of some sea snakes. Body form of swimming snakes differed significantly from their shape when resting on a flat surface or when anesthetized and suspended in water. The extent of lateral flattening is positively correlated with swimming speed, a relationship generally supported by tests of trunk models in a flow tank. In Nerodia, the ability to temporarily flatten the trunk depends on kinetic costovertebral joints, a large compressible body cavity, and the absence of ventral skeletal support - features found in most snakes. Histological studies and manipulations of partially dissected preserved specimens showed that the resting angle of the ribs is maintained by localized elastic hypertrophy of the costovertebral capsular ligament. Trunk form during swimming in Nerodia is proposed to arise from anteromedial movement of the distal rib powered by deep muscles acting in concert with those proposed to generate undulation of the vertebral column.     

Effects of temperature and perch diameter on arboreal locomotion in the snake Elaphe guttata

Arboreality is widespread in multiple lineages of snakes and these habitats are important for foraging, escaping predators, and thermoregulation for many species. However, very little is known about factors influencing the arboreal locomotor abilities of snakes. Arboreal performance was assessed in a semi-arboreal snake (Elaphe guttata) using an artificial perch apparatus. Locomotor velocity, body posture, and balance was measured during movement on three perch diameters (3, 6, 10-cm) at three temperatures (10, 20, 30 degrees C). Velocities attained by E. guttata on perches are much slower than those of terrestrial lateral undulation and swimming and somewhat slower than concertina velocities recorded in other species across the same experimental temperatures. At higher temperatures, faster speeds were associated with a more elongated posture. At lower temperatures, snakes displayed a more looped body posture, but still fell more often than at higher temperatures. Our results suggest that temperature has a large influence on both balance and movement by snakes on perches. Although there were no differences in velocities resulting from perch diameter, snakes fell more often from thicker perches. This differs from arboreal velocities attained by limbed vertebrates, which decrease with decreasing perch diameter, suggesting that snakes have a size-relative advantage over limbed animals, such as lizards, when traversing a network of narrow branches. Future studies investigating arboreal locomotion among snakes that vary both phylogenetically and morphologically are needed to assess the potential benefits of limblessness in complex, three-dimensional environments.     

Habitat use and movement patterns of the viviparous aquatic snake, Oocatochus rufodorsatus, from northeast Asia

To determine the habitat usage and movement patterns of the viviparous aquatic snake Oocatochus rufodorsatus (formerly Elaphe rufodorsata), we radio-tracked 21 snakes on agricultural lands during two active seasons in 2007 and 2008. Male and female snakes stayed close to aquatic habitats such as paddy fields and agricultural ponds during both breeding and non-breeding periods, except when the snakes moved to dry terrestrial areas to hibernate in late fall. The use of different structural features in the habitat, such as ground, tree, underground, and water, varied depending on the air and water temperatures, female's reproductive conditions, and the time of day. Male and female snakes moved about 17 m daily and postpartum females moved farther than antepartum females. The home ranges of males and females were 0.45 ha and 0.47 ha, respectively, and the year-round home range of this species was approximately 1.54 ha (95% fixed kernel). Thus, to conserve a population of O. rufodorsatus in our study area, areas including both aquatic and terrestrial habitats within a radius of 150 m from a core pond habitat must be preserved.     

Red and white muscle activity and kinematics of the escape response of the bluegill sunfish during swimming

Summary We quantified midline kinematics with synchronized electromyograms (emgs) from the red and white muscles on both sides of bluegill sunfish (Lepomis macrochirus) during escape behaviors which were elicited from fish both at a standstill and during steady speed swimming. Analyses of variance determined whether or not kinematic and emg variables differed significantly between muscle fiber types, among longitudinal positions, and between swimming versus standstill trials.At a given longitudinal location, both the red and white muscle were usually activated synchronously during both stages of the escape behavior. Stage 1 emg onsets were synchronous; however, the mean durations of stage 1 emgs showed a significant increase posteriorly from about 11 to 15 ms. Stage 2 emgs had significant posterior propagation, but the duration of the stage 2 emgs was constant (17 ms). Posterior emgs from both stages occurred during lengthening of the contractile tissue (as indicated by lateral bending). Steady swimming activity was confined to red muscle bursts which were propagated posteriorly and had significant posterior decrease in duration from about 50% to 37% of a cycle. Fish performed escape responses during all phases of the steady swimming motor pattern. All kinematic events were propagated posteriorly. Furthermore, no distinct kinematic event corresponded to the time intervals of the stage 1 and 2 emgs. The rate of propagation of kinematic events was always slower than that of the muscle activity. The phase relationship between lateral displacement and lateral bending also changed along the length of the fish. Escape responses performed during swimming averaged smaller amplitudes of stage 2 posterior lateral displacement; however, most other kinematic and emg variables did not vary significantly between these two treatments.Abbreviations A angle of lateral flexion (bending) of midline at a single point in time - A1, A2 change in A from T₀ to T₁ and from T₁ to T₂ - AMX maximal lateral flexion concave towards the side of the stage 1 emg - AMXR equals AMX minus A at T₀ - AT1, AT2 lateral flexion at T₁ and T₂ - DUR1, DUR2 durations of stage 1 and stage 2 emgs - emg electromyogram - ON2 onset time of stage 2 emg - RELDUR relative duration of steady swimming emg - T₀, T₁, T₂ times of stage 1 emg onset, latest stage 1 emg offset and latest stage 2 emg offset standardized such that T₀ = 0 - TAMX, TAMN, TYMX times of maximal lateral flexion, no lateral flexion and maximum lateral displacement - Y1, Y2 amounts of lateral displacement from T₀ to T₁ and from T₁ to T₂ - YMXR relative amount of lateral displacement from T₀ to TYMX     

CONSTRAINTS ON REPRODUCTIVE INVESTMENT: A COMPARISON BETWEEN AQUATIC AND TERRESTRIAL SNAKES

Life-history theory predicts that “costs” of reproduction may be important evolutionary determinants of reproductive investment; previous studies on reptiles indicate that decrements to maternal mobility may be among the most important components of such costs. Biomechanical models suggest that reproductive investment in aquatic snakes may be constrained by the important locomotory role of the posterior part of the body during swimming: carrying eggs or offspring in this region would more seriously impair locomotory efficiency in swimming than in terrestrial lateral undulation. If this constraint is important, aquatic snakes would be expected to have lower clutch masses relative to body mass than terrestrial species and to carry the clutch in a more anterior position (commencing at the same proportion of maternal body length anteriorly, but not extending as far posteriorly). Comparisons between aquatic and terrestrial snakes of several families confirm these predictions. Phylogenetic analysis suggests that this pattern of reduced reproductive investment has evolved independently in each of the four ophidian lineages that contain marine species (acrochordids, homalopsine colubrids, laticaudid sea snakes, and hydrophiid sea snakes). Although it thus seems likely that these patterns represent adaptations to aquatic versus terrestrial life, the nature of the selective forces involved remains speculative. The hypothesis based on locomotory impairment of gravid females has better empirical support than any alternative hypothesis, as it successfully predicts modifications in the position of the clutch within the female's body, as well as overall reduced reproductive investment.     

Morphological and biochemical evidence for the evolution of salt glands in snakes

Vertebrate salt glands have evolved independently multiple times, yet there are few hypotheses about the processes underlying the convergent evolution of salt glands across taxa. Here, we compare the morphology and molecular biology of specialized salt-secreting glands from a marine snake (Laticauda semifasciata) with the cephalic glands from semi-marine (Nerodia clarkii clarkii) and freshwater (N. fasciata) watersnakes to look for evidence of a salt gland in the former and to develop hypotheses about the evolution of snake salt glands. Like the salt gland of L. semifasciata, the nasal and anterior/posterior sublingual glands in both species of Nerodia exhibit a compound tubular shape, and express basolateral Na(+)/K(+)-ATPase (NKA) and Na(+)/K(+)/2Cl(-)cotransporter (NKCC); however, the abundance of NKA and NKCC in N. fasciata appears lower than in N. c. clarkii. Aquaporin 3 (AQP3) is also basolateral in the sublingual glands of both species of Nerodia, as is abundant neutral mucin; both AQP3 and mucin are absent from the salt gland in L. semifasciata. Thus, we propose that the evolution of the snake salt gland by co-option of an existing gland involved at least two steps: (i) an increase in the abundance of NKA and NKCC in the basolateral membranes of the secretory epithelia, and (ii) loss of AQP3/mucus secretion from these epithelia.     

Adrenergic innervation of the large arteries and veins of the semiarboreal rat snake Elaphe obsoleta

Fluorescence histochemistry was used to study the adrenergic innervation of the large arteries and veins at six points along the body of the semiarboreal rat snake Elaphe obsoleta. Apart from the vessels adjacent to the heart, there was a marked contrast in the density of adrenergic innervation of anterior and posterior systemic arteries and veins. The anterior arteries and veins have little adrenergic innervation in contrast to the extremely dense innervation of the arteries and veins posterior to the heart. The innervation pattern is consistent with known physiological adjustments to gravity and suggests a mechanism for regulating dependent blood flow via sympathetic nerves. In comparison to the posterior systemic arteries, parallel segments of pulmonary artery taken from the same body position of Elaphe contained a much sparser innervation by adrenergic nerves. The sparser innervation can be correlated with less gravitational disturbance in the pulmonary artery, which is relatively short in this and in other arboreal snakes.     

Fiber‐type distribution of the perivertebral musculature in Ambystoma

Many salamanders locomote in aquatic and terrestrial environments. During swimming, body propulsion is solely produced by the axial musculature generating lateral undulations of the trunk and tail. During terrestrial locomotion, the trunk is oscillated laterally in a standing wave, and body propulsion is achieved by concerted trunk and limb muscle action. The goal of this study was to increase our knowledge of the functional morphology of the tetrapod trunk. We investigated the muscle‐fiber‐type distribution and the anatomical cross‐sectional area of all perivertebral muscles in Ambystoma tigrinum and A. maculatum. Muscle‐fiber‐type composition was determined in serial cross‐sections based on m‐ATPase activity. Five different body segments were investigated to test for cranio‐caudal changes along the trunk. The overall fiber‐type distribution was very similar between the species, but A. tigrinum had relatively larger muscles than A. maculatum, which may be related to its digging behavior. None of the perivertebral muscles possessed a homogeneous fiber‐type composition. The M. interspinalis showed a distinct layered organization and may function to ensure the integrity of the spine (local stabilization). The M. dorsalis trunci exhibited the plesiomorphic pattern for notochordates in having a distinct superficial layer of red and intermediate fibers, which covered the central white fibers; therefore, it is suggested to function as a mobilizer and a stabilizer of the trunk, but, may also be involved in modulating body stiffness. Similarly, the M. subvertebralis showed clear regionalizations, implying functional subunits that can stabilize and mobilize the trunk as well as modulate of body stiffness. Cranio‐caudally, neither the fiber‐type composition nor the a‐csa changed dramatically, possibly reflecting the need to perform well in both aquatic and terrestrial habitats. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

发酵床养蛇初探

目的探讨发酵床模式养殖尖吻蝮、舟山眼镜蛇和王锦蛇的可行性。方法分别选取3种健康驯饲好的4月龄蛇苗,各分成2组,一组采用传统沙床饲养模式,另一组采用发酵床饲养模式,饲养8个月后,通过比较蛇的体重、料肉比及环境氨气浓度变化,分析发酵床养殖模式优劣。结果就尖吻蝮养殖效果而言,采用两种养殖模式无显著性差异;但采用发酵床模式对眼镜蛇和王锦蛇饲养相比沙床模式具有明显优势,表现为增重更快、料肉比更小,且氨气浓度显著降低;氨气浓度及体重变化分析结果表明,5ppm以上浓度对王锦蛇生长有一定影响,10ppm以上浓度对眼镜蛇生长有一定影响。结论发酵床技术可用于蛇类养殖,而且总体上优于传统沙床模式;此外,发酵床模式尤其适宜如眼镜蛇和王锦蛇等活动较多、进食量较大、排便量大的蛇类养殖。     

Haemogregarine specificity in two communities of Florida snakes, with descriptions of six new species of Hepatozoon (Apicomplexa: Hepatozoidae) and a possible species of Haemogregarina (Apicomplexa: Haemogregarinidae)

Five species of snakes in Florida, from Palm Beach County in the south and Alachua County 450 km to the north, occur in similar habitat but have distinctive Hepatozoon species characteristic of each host species. In Palm Beach County, Diadophis punctatus is host to Hepatozoon punctatus n. sp., Thamnophis sauritus sackenii to Hepatozoon sauritus n. sp., and Nerodia fasciata pictiventris to Hepatozoon pictiventris n. sp. In Alachua County, N. fasciata pictiventris is parasitized by Hepatozoon fasciatae n. sp., Seminatrix p. pygaea by Hepatozoon seminatrici n. sp., and Thamnophis s. sirtalis by Hepatozoon sirtalis n. sp. Each Hepatozoon sp. has distinctive gamonts and sporogonic characters and, in the 4 species where known, meronts. Nerodia floridana is host to Haemogregarina floridana n. sp. in both localities, with generic identification tentative, based upon presence of erythrocytic meronts. The presence of sporocysts in the proboscis of 31% of Aedes aegypti infected by H. pictiventris is the first report of infective stages of a reptilian Hepatozoon species within the mouthparts of a dipteran vector. This study suggests that in Florida, at least, the diversity of the Hepatozoon community not only equals but probably exceeds the diversity of the snake communities present, and that host specificity in nature may be much greater than that postulated from previous studies.     

Alpha-Glycerophosphate dehydrogenase (insoluble) and lactic dehydrogenase activities in the skeletal muscles of two insects.

The flight muscle preparations of the dragonfly Pantala flavescens and the aquatic beetle Cybister confusus showed extremely low levels of lactic dehydrogenase activity and high levels of alpha-glycerophosphate dehydrogenase (insoluble) activity. The activities of these two enzymes in the leg muscle of the beetle were approximately the same (1:1), but lactic dehydrogenase activity was several times higher than that in the flight muscles of both Insects. These results have been interpreted as indicating the high energy-yielding demands of the flight muscles during continuous sustained activity, while the leg muscles of the beetle which are involved in swimming activity derive their energy predominantly through anaerobic glycolysis.     

Identification of beta-type phospholipase A(2) inhibitor in a nonvenomous snake,Elaphe quadrivirgata

A novel serum protein inhibiting specifically the enzymatic activity of the basic phospholipase A(2) (PLA(2)) from the venom of the Chinese mamushi snake (Agkistrodon blomhoffii siniticus) was purified from a nonvenomous Colubridae snake, Elaphe quadrivirgata. The purified inhibitor was a 150-kDa glycoprotein having a trimeric structure, composed of two homologous 50-kDa subunits. Their amino acid sequences, containing leucine-rich repeats, were typical of the beta-type PLA(2) inhibitor (PLIbeta), previously identified from the serum of A. blomhoffii siniticus. The inhibitor inhibited exclusively group II basic PLA(2)s and did not inhibit other kinds of PLA(2)s. This is the first paper reporting the existence of PLIbeta in a nonvenomous snake. The existence of PLIbeta in the nonvenomous snake reflects that PLIbetas are widely distributed over the snake species and participate commonly in regulating the physiological activities of the unidentified target PLA(2)s.     

A comparative study of locomotion in the caecilians Dermophis mexicanus and Typhlonectes natans (Amphibia: Gymnophiona)

We compared locomotion of two species of caecilian using x-ray videography of the animals traversing smooth-sided channels and a pegboard. Two channel widths were used, a body width channel and a body width + 20% channel. The terrestrial caecilian, Dermophis mexicanus , used internal concertina locomotion in both channels and lateral undulation on the pegboard. The aquatic caecilian, Typhlonectes natans , was not able to move at all in the body width channel. In the wider channel Typhlonectes proceeded at the same speed as Dermophis while using normal, rather than internal, concertina locomotion. On the pegboard, Typhlonectes used lateral undulation and achieved 2.5 times the speed managed by Dermophis. A phylogenetic analysis of this, and other, evidence shows that (1) internal concertina evolved in the ancestor to extant caecilians and (2) internal concertina locomotion was secondarily lost in the aquatic caecilians.     

Red muscle function in stiff-bodied swimmers: there and almost back again

Fishes with internalized and endothermic red muscles (i.e. tunas and lamnid sharks) are known for a stiff-bodied form of undulatory swimming, based on unique muscle-tendon architecture that limits lateral undulation to the tail region even though the red muscle is shifted anteriorly. A strong convergence between lamnid sharks and tunas in these features suggests that thunniform swimming might be evolutionarily tied to this specialization of red muscle, but recent observations on the common thresher shark (Alopias vulpinus) do not support this view. Here, we review the fundamental features of the locomotor systems in lamnids and tunas, and present data on in vivo muscle function and swimming mechanics in thresher sharks. These results suggest that the presence of endothermic and internalized red muscles alone in a fish does not predict or constrain the swimming mode to be thunniform and, indeed, that the benefits of this type of muscle may vary greatly as a consequence of body size.     

Functional differences between anatomical regions of the anconeus muscle in humans

This study sought to resolve a longstanding debate of the function of anconeus. Intramuscular and surface electromyography electrodes recorded muscle activity from two regions of anconeus and from typical elbow flexion and extension muscles. Eleven participants performed pronation–supination around the medial and lateral axes of the forearm, elbow flexion–extension in pronation, supination and neutral positions of the forearm, and gripping. Maximal voluntary contractions (MVC) and submaximal (10% MVC) force-matching tasks were completed. Activity varied between longitudinal (AL) and transverse (AT) segments of anconeus. Although both muscle regions were active across multiple directions (including opposing directions), AL was more active during pronation than supination, whereas AT showed no such difference. During pronation, activity of AL and AT was greatest about the lateral forearm axis. AT was more active during elbow extension with the forearm in pronation, whereas AL did not differ between pronated and neutral forearm alignment. These findings are consistent with the proposal that AL makes a contribution to control of abduction of the ulna during forearm pronation. Different effects of forearm position on AL and AT activity during elbow extension may be explained by the anatomical differences between the regions. These data suggest anconeus performs multiple functions at the elbow and forearm and this varies between anatomically distinct regions of the muscle.     

The role of hind limb flexor muscles during swimming in the toad, Bufo marinus

Most work examining muscle function during anuran locomotion has focused largely on the roles of major hind limb extensors during jumping and swimming. Nevertheless, the recovery phase of anuran locomotion likely plays a critical role in locomotor performance, especially in the aquatic environment, where flexing limbs can increase drag on the swimming animal. In this study, I use kinematic and electromyographic analyses to explore the roles of four anatomical flexor muscles in the hind limb of Bufo marinus during swimming: m. iliacus externus, a hip flexor; mm. iliofibularis and semitendinosus, knee flexors; and m. tibialis anticus longus, an ankle flexor. Two general questions are addressed: (1) What role, if any, do these flexors play during limb extension? and (2) How do limb flexors control limb flexion? Musculus iliacus externus exhibits a large burst of EMG activity early in limb extension and shows low levels of activity during recovery. Both m. iliofibularis and m. semitendinosus are biphasically active, with relatively short but intense bursts during limb extension followed by longer and typically weaker secondary bursts during recovery. Musculus tibialis anticus longus becomes active mid way through recovery and remains active through the start of extension in the next stroke. In conclusion, flexors at all three joints exhibit some activity during limb extension, indicating that they play a role in mediating limb movements during propulsion. Further, recovery is controlled by a complex pattern of flexor activation timing, but muscle intensities are generally lower, suggesting relatively low force requirements during this phase of swimming.     

Locomotion pattern and trunk musculoskeletal architecture among Urodela

We comparatively examined the trunk musculature and prezygapophyseal angle of mid‐trunk vertebra in eight urodele species with different locomotive modes (aquatic Siren intermedia, Amphiuma tridactylum, Necturus maculosus and Andrias japonicus; semi‐aquatic Cynops pyrrhogaster, Cynops ensicauda; and terrestrial Hynobius nigrescens, Hynobius lichenatus and Ambystoma tigrinum). We found that the more terrestrial species were characterized by larger dorsal and abdominal muscle weight ratios compared with those of the more aquatic species, whereas muscle ratios of the lateral hypaxial musculature were larger in the more aquatic species. The lateral hypaxial muscles were thicker in the more aquatic species, whereas the M. rectus abdominis was more differentiated in the more terrestrial species. Our results suggest that larger lateral hypaxial muscles function for lateral bending during underwater locomotion in aquatic species. Larger dorsalis and abdominal muscles facilitate resistance against sagittal extension of the trunk, stabilization and support of the ventral contour line against gravity in terrestrial species. The more aquatic species possessed a more horizontal prezygapophyseal angle for more flexible lateral locomotion. In contrast, the more terrestrial species have an increasingly vertical prezygapophyseal angle to provide stronger column support against gravity. Thus, we conclude trunk structure in urodeles differs clearly according to their locomotive modes.     

Elasmosaur (Reptilia: Sauropterygia) neck flexibility: implications for feeding strategies

Elasmosaurs were extremely long-necked, aquatic reptiles that used four flippers for locomotion. Their distinctive long neck distinguishes them from all other Mesozoic forms, yet the potential uses and constraints of this structure are poorly understood, particularly with regard to feeding. Several associated series of elasmosaurian cervical vertebrae were used to measure ranges of potential flexion. Two-dimensional models, based on a complete specimen of the Late Cretaceous elasmosaur Aphrosaurus furlongi, were created to measure mobility in both vertical and horizontal planes. Accuracy of the models was assessed through comparative analyses with currently extant vertebrate analogues (e.g. snake, turtle, seal). Results suggest that the elasmosaurian neck was capable of a 75-177 degrees ventral, 87-155 degrees dorsal, and 94-176 degrees lateral range of movement depending upon the thickness of cartilage reconstructed between each vertebra. Neck postures such as a 'swan-like' S-shape are shown to be implausible because they require >360 degrees vertical flexion. However, maintenance of a straight neck while swimming, together with considerable lateral and/or ventral movement during prey capture and feeding are feasible.     

Elasmosaur (Reptilia: Sauropterygia) neck flexibility: implications for feeding strategies.

Elasmosaurs were extremely long-necked, aquatic reptiles that used four flippers for locomotion. Their distinctive long neck distinguishes them from all other Mesozoic forms, yet the potential uses and constraints of this structure are poorly understood, particularly with regard to feeding. Several associated series of elasmosaurian cervical vertebrae were used to measure ranges of potential flexion. Two-dimensional models, based on a complete specimen of the Late Cretaceous elasmosaur Aphrosaurus furlongi, were created to measure mobility in both vertical and horizontal planes. Accuracy of the models was assessed through comparative analyses with currently extant vertebrate analogues (e.g. snake, turtle, seal). Results suggest that the elasmosaurian neck was capable of a 75-177 degrees ventral, 87-155 degrees dorsal, and 94-176 degrees lateral range of movement depending upon the thickness of cartilage reconstructed between each vertebra. Neck postures such as a 'swan-like' S-shape are shown to be implausible because they require >360 degrees vertical flexion. However, maintenance of a straight neck while swimming, together with considerable lateral and/or ventral movement during prey capture and feeding are feasible.