Satellite tracking and diving behaviour of sub-adult narwhals (<Emphasis Type="Italic">Monodon monoceros</Emphasis>) in Svalbard,Norway

Three juvenile narwhals captured during August 1998 in the northeast of Svalbard, Norway, were equipped with satellite-relayed data loggers (SRDLs) that transmitted diving and swim-speed data, in addition to location, for up to 46 days. A total of 1,354 complete dive cycles were recorded. Most of the diving was shallow and of short duration. Maximum recorded dive depth was 546 m, maximum recorded dive duration was 24.8 min, and maximum recorded swim-speed was 4.7 ms⁻¹. Ascent speed, vertical ascent speed, descent speed and vertical descent speed were all significantly higher during deep dives (>200 m) than for shallow dives (<200 m). In addition both ascent and descent angles were much steeper for deep dives than during shallow dives. Most of the shallow diving seemed to be associated with travelling, with the animal shifting between various locations, while the deep diving (often to the bottom) for extended periods in some specific areas might have been associated with foraging. Even though the sample size in this study is small, the data are the first information available for movements and diving behaviour of narwhals near Svalbard.     

Costs of diving by wing and foot propulsion in a sea duck,the white-winged scoter

Most birds swim underwater by either feet alone or wings alone, but some sea ducks often use both. For white-winged scoters (Melanitta fusca), we measured costs of dives to 2 m with descent by feet only versus wings + feet (only feet are used at the bottom). Dive costs repaid during the recovery period after a dive bout were an important fraction (27–44%) of total dive costs, and removing costs of extraneous surface behaviors increased resolution of differences between dive types. Scoters using wings + feet had 13% shorter descent duration, 18% faster descent speed, 31% fewer strokes/m, and 59% longer bottom duration than with feet only. The cost of time underwater for dives using wings + feet was 32–37% lower than with feet only (P = 0.09 to 0.15). When indirect methods were used to partition descent costs from costs of ascent and bottom phases, using wings + feet lowered descent cost by an estimated 34%. Thus, using wings + feet increases descent speed and lowers descent cost, leaving more time and energy for bottom foraging. For birds in cold water, the large savings may result from both biomechanical and thermoregulatory factors.     

Discrete movements of foot epithelium during adhesive locomotion of a land snail

During the adhesive locomotion of land snails a series of short dark transverse bands, called pedal or foot waves, is visible ifa moving snail's ventral surface is observed through a sheet of glass. Moreover, the mucus secreted from the pedal glands and some pedal epithelial cells forms a thin layer which acts as a glue augmenting adherence, while also acting as a lubricant under the moving parts of the snail's foot. The relationships between velocity and the frequency of pedal waves as well as changes in the volume of small air bubbles under foot waves were analyzed by means of digital recordings made through a glass sheet on which the snails were moving. On the ventral surface of a moving snail foot, the adhering parts of the foot constituted about 80% of the total area, while several moving parts only about 20%. The single moving region of the foot (the pedal wave) amounted to about 3% of snail length. The epithelium in the region of the pedal wave was arched above the substrate and was also more wrinkled than the stationary epithelium, which enabled the forward motion of each specific point of epithelium during the passage of a pedal wave above it. The actual area of epithelium engaged by a pedal wave was at least 30% greater than the area of the epithelium as recorded through a glass sheet. In the region of the pedal wave, the tiny subepithelial muscles acting on the epithelium move it up in the front part of the wave, and then down at the end of the wave, operating vertically in relation to the substrate. In the middle part of the wave, the epithelium only moves forward. In summary, during the adhesive locomotion of snails, the horizontal movement of the ventral surface epithelium proceeds as temporally separate phases of upward, forward and downward movement.     

Respiratory frequency, dive behaviour and social interactions of leatherback turtles, Dermochelys coriacea during the inter-nesting interval

We collected simultaneous dive Time Depth Recorder (TDR) data and video images from free swimming adult female leatherback turtles, Dermochelys coriacea, during the first 24 h after nesting on the beach, in order to determine relationships between dive parameters, activity, overall respiratory frequency and behaviour.We identified three different underwater locomotory activities (subsurface swimming, V-shaped dives and U-shaped dives) from video and TDR data that varied in their mean depth, duration and a number of other parameters. Overall respiratory frequency (overall f_R) was significantly different between all locomotory activities, with turtles taking 1.7±0.1 breaths min⁻¹ while subsurface swimming, 0.78 breaths min⁻¹ after V-shaped dives and 0.57 breaths min⁻¹ after U-shaped dives. Descent rates and ascent rates were significantly faster in U-shaped dives (descent 0.19±0.010 m s⁻¹, ascent 0.28±0.015 m s⁻¹) than in V-shaped dives (descent 0.10±0.008 m s⁻¹, ascent 0.12±0.012 m s⁻¹). Flipper stroke rates were significantly lower during the bottom component of U-shaped dives (0.18±0.02 strokes s⁻¹) than during the descent (0.29±0.03 strokes s⁻¹) or ascent (0.29±0.03 strokes s⁻¹). From overall f_R and flipper stroke rate data, we inferred that turtles used less energy during U-shaped dives than the other activity types. We recorded interactions between male turtles and the study females that lasted up to 11 min, during which male turtles displayed the characteristic courtship behaviour of sea turtles. It appeared that females attempted to avoid males by aborting ascent and extending dive duration to swim to the sea floor when males were present.     

Vertical movement and time allocation of a freshwater pulmonate snail

The paper presents an analysis of locomotion and location in a vertical water column of 10Biomphalaria glabrata, under constant conditions of light, temperature, and food availability. Individual snails varied in distance traveled, and in the percentage of time spent in different areas of the water column. Distance covered ranged from 53 cm to 100 cm h^–1 . Approximately 21% of locomotion was observed to be passive (floating up or sinking down). Snails spent approximately 58% of the time in the bottom section, 35% in the top section and only 7% in the middle region. The study provides baseline data that can be used to compare further studies ofBiomphalaria glabrata as various conditions are altered.     

Locomotion in the pulmonate snail Melampus--II. Recovery after pedal ganglion excision

1. When one pedal ganglion is removed, snails first crawl using the unoperated side of the foot, but in 4-8 weeks the operated side exhibits an anterior-to-posterior gradient of recovery. 2. A ganglion bud bridges the site of the missing ganglion and axons project from intact central ganglia into the foot. 3. Rhythmic activity in right and left pedal nerve pairs is correlated during locomotion in the regenerated snails. 4. The oscillator in the remaining pedal ganglion drives bilaterally coordinated activity. Regenerated projections from the cerebral ganglia through the bud to the remaining pedal ganglion suffice to initiate locomotion.     

Three-dimensional movements and swimming activity of a northern elephant seal

We attached a video system and data recorder to a northern elephant seal to track its three-dimensional movements and observe propulsive strokes of the hind flippers. During 6 h of recording, the seal made 20 dives and spent 90% of the time submerged. Average dive duration, maximum depth and swimming speed were 14.9 min+/-6.1 S.D., 289 m+/-117 S.D. and 1.1 m s(-1)+/-0.12 S.D., respectively. The distance swum during a dive averaged 925 m+/-339 S.D., and the average descent and ascent angles were 41 degrees +/-18 S.D. and 50 degrees +/-21 S.D., respectively. Dive paths were remarkably straight suggesting that the seal was navigating while submerged. We identified three modes of swimming based on the interval between propulsive strokes: continuous stroking; stroke-and-glide swimming; and prolonged gliding. The seal used continuous stroking from the surface to a mean depth of 20 m followed by stroke-and-glide swimming. Prolonged gliding started at a mean depth of 60 m and continued to the bottom of dives. For dives to depths of 300 m or more, 75% of the descent time was spent in prolonged gliding and 10% in stroke-and-glide swimming, amounting to 5.9-9.6 min of passive descent per dive. Average swimming speed varied little with swimming mode and was not a good indicator of propulsive effort. It appears that the seal can use prolonged gliding to reduce the cost of transport and increase dive duration. Energetically efficient locomotion may help explain the long and deep dives that routinely exceed the theoretical aerobic dive limit in this species.     

GEOTROPISM AND MUSCLE TENSION IN HELIX

1. The snail Helix aspersa Müller, is negatively geotropic during the daytime, but positive or indifferent at night. 2. The precision of geotropic orientation is a function of the gravity component acting on the body. 3. The rate of geotropic locomotion is also determined by the gravity component (sine of the angle of inclination). 4. The rate of upward movement is increased 1.51 times at 45° inclination by loading the snail with one-half its weight. No such increase is seen in loaded snails creeping on a horizontal surface. 5. Moderate centrifugation results in orientation and locomotion towards the center of rotation. 6. A response analogous to the homostrophic reflex occurs when a backward pull to right or to left is exerted on the shell. Bilaterally equal tension applied to the shell causes locomotion along a path parallel and opposite to the direction of the pull. 7. All the observations go to show that the stimulus for geotropic orientation and locomotion is tension of the body muscles produced by the downward pull of gravity, and that the stimulus is received by the proprioreceptors of these muscles. Otolith apparatus and analogous organs, when present, may assist in the response, but they do not seem to be requisite in all cases. Since the precision of orientation and the rate of locomotion are functions of the gravity component acting on the body, the muscle tension theory of the geotropic reactions accords fully with Loeb''s tropism doctrine for animals.     

Locomotion in diving elephant seals: physical and physiological constraints

To better understand how elephant seals (Mirounga angustirostris) use negative buoyancy to reduce energy metabolism and prolong dive duration, we modelled the energetic cost of transit and deep foraging dives in an elephant seal. A numerical integration technique was used to model the effects of swim speed, descent and ascent angles, and modes of locomotion (i.e. stroking and gliding) on diving metabolic rate, aerobic dive limit, vertical displacement (maximum dive depth) and horizontal displacement (maximum horizontal distance along a straight line between the beginning and end locations of the dive) for aerobic transit and foraging dives. Realistic values of the various parameters were taken from previous experimental data. Our results indicate that there is little energetic advantage to transit dives with gliding descent compared with horizontal swimming beneath the surface. Other factors such as feeding and predator avoidance may favour diving to depth during migration. Gliding descent showed variable energy savings for foraging dives. Deep mid-water foraging dives showed the greatest energy savings (approx. 18%) as a result of gliding during descent. In contrast, flat-bottom foraging dives with horizontal swimming at a depth of 400m showed less of an energetic advantage with gliding descent, primarily because more of the dive involved stroking. Additional data are needed before the advantages of gliding descent can be fully understood for male and female elephant seals of different age and body composition. This type of data will require animal-borne instruments that can record the behaviour, three-dimensional movements and locomotory performance of free-ranging animals at depth.     

Creeping patterns of human adults and infants

The patterns of swing and support for the hands-and-feet or hands-and-knees gaits (creeping) of human adults and infants are compared based on data from a number of studies. Human infants show considerable variability in their creeping gait patterns, whereas adult patterns are less variable and fairly consistent after a few minutes of practice. Creeping on hands-and-knees appears to dictate a gait pattern characteristically different from creeping on hands-and-feet. The highly inefficient nature of adult creeping supports the view that our early hominid ancestors were poorly adapted to quadrupedal locomotion. Data obtained from high-speed film analysis of human creeping patterns show that the number of foot lengths per stride in creeping is about twice that for normal upright walking at the same speed. The support pattern of human creeping is different from that of nonhuman primates. These findings are discussed in the context of debate concerning the origin of the Laetoli hominid footprints and the knuckle-walking hypothesis.     

Dive form and function in belugas Delphinapterus leucas of the eastern Canadian High Arctic

The underwater behaviour of 11 belugas or white whales was examined during summer using time-at-depth records relayed by satellite-linked data-loggers. Simultaneous tracking information was obtained for each whale. Eight distinct dive profiles were identified in submergences made to depths of >40 m. Four of these, together comprising 84% of these “deep” dives, were of a square profile. They were characterised by a continuous descent to a particular depth (usually the sea bed), a “bottom phase” at or near that depth, and a direct ascent to the surface. These dives are presumed to be made for benthic foraging. Other, much less common, dive shapes were “V”-shaped, parabolic and trapezoidal. “Shallow” dives (15–40 m depth) were of a variety of shapes, short duration and high average horizontal speeds. Many probably occurred during periods of directed travel. This population of belugas treats most of the water column as dead space separating resources of oxygen and nutrition. Received: 8 January 1998 / Accepted: 27 April 1998     

Morpho-functional modifications of human syncytiotrophoblast plasma membrane during Pregnancy Induced Hypertension

Changes in [³⁵S]methionine protein labeling patterns were examined by following incorporation into the acid precipitate protein fraction of land snails,Otala lactea (Müller) (Pulmonata, Helicidae). Labeled proteins were analyzed by SDS polyacrylamide gel electrophoresis and isoelectric focusing columns. Snails in four different physiological states were compared: active controls, short term aestivating snails (injected and allowed to enter aestivation), long term aestivating snails (aestivated for 14 days, injected, and maintained in the aestivating state), and snails aroused after aestivation (aestivated, injected, and aroused). Protein associated radioactivity was measured over a 7 day time course post injection. Autoradiographic analysis of SDS-polyacrylamide gels showed increases in the radioactivity of four proteins: 91 kDa (hepatopancreas, day 1 in long term aestivating animals), 50 kDa (hepatopancreas, day 2 in short term aestivating snails), 70 kDa and 30 kDa (foot, day 2 in short term aestivating animals). Hepatopancreas and foot from day 1 long term aestivating and day 2 short term aestivating animals were also analyzed by isoelectric focusing columns. Several pH-specific differences were apparent when controls and aestivating animals were analyzed. In particular a peak of radioactivity was observed at pH 5.05 in 1 d long term aestivating hepatopancreas and at pH 4.30 in 2d short term aestivating animals. Several differences were noted in foot with no specific pattern emerging. SDS-polyacrylamide gel electrophoresis analysis of the hepatopancreas peaks showed the appearance of several bands with increased radioactivity, including the 91 kDa and 50 kDa proteins described above. These results suggest thatO. lactea aestivation specific proteins may be involved in the transition to a depressed metabolic state.Abbreviations dpm radioactive disintegrations per minute - PAGE polyacrylamide gel electrophoresis - SDS sodium dodecyl sulphate - SRP stress related protein     

Functional morphology of the muscles in Philodina sp. (Rotifera: Bdelloidea)

Whole-mounts of Philodina sp., a bdelloid rotifer, were stained with fluorescent-labeled phalloidin to visualize the musculature. Several different muscle types were identified including incomplete circular bands, coronal retractors and foot retractors. Based on the position of the larger muscle bands in the body wall, their function during creeping locomotion and tun formation was inferred. Bdelloid creeping begins with the contraction of incomplete circular muscle bands against the hydrostatic pseudocoel, resulting in an anterior elongation of the body. One or more sets of ventral longitudinal muscles then contract bringing the rostrum into contact with the substrate, where it presumably attaches via adhesive glands. Different sets of ventral longitudinal muscles, foot and trunk retractors, function to pull the body forward. These same longitudinal muscle sets are also used in `tun' formation, in which the head and foot are withdrawn into the body. Three sets of longitudinal muscles supply the head region (anterior head segments) and function in withdrawal of the corona and rostrum. Two additional pairs of longitudinal muscles function to retract the anterior trunk segments immediately behind the head, and approximately five sets of longitudinal retractors are involved in the withdrawal of the foot and posterior toes. To achieve a greater understanding of rotifer behavior, it is important to elucidate the structural complexity of body wall muscles in rotifers. The utility of fluorescently-labeled phalloidin for the visualization of these muscles is discussed and placed in the context of rotifer functional morphology.     

Voluntary bipedal walking of infant chimpanzees

The voluntary bipedal walking of infant chimpanzees was studied by the analysis of foot force and by motion analysis. The infants were trained to locomote on a level platform without any restrictions on the locomotor pattern. The voluntary bipedal walking was compared with the other types of locomotion at the same age and with the trained bipedal walking performed by other chimpanzees, including adult chimpanzees. The characteristics of voluntary bipedal walking in the infant until one year of age were: (1) high-speed walking with short cycle duration; (2) short stance phase duration; (3) small braking component of the preceding leg and large acceleration of the following leg; (4) one downward peak in the vertical component; and (5) a relatively small transverse component. Bipedal walking usually continued for less than one second and ended in quadrupedal locomotion. During walking, the preceding foot touched the floor, heel first, as in the case of older chimpanzees and humans. At this age, bipedal walking was similar to high-speed locomotion. The voluntary bipedal walking of the two-year-old and frour-yearold chimpanzees was characterized as follows: (1) slower speed than during quadrupedal locomotion, (2) relatively long periods and distances; (3) well balanced accelerating and braking components; and (4) a vertical component showing two downward peaks and a trough in between during numerous trials. The last characteristic means that the body center of gravity is higher in the single stance phase, just as in the bipedal walkinbg of the adult chimpanzees and humans. The bipedal walking of infant chimpanzees was discussed in comparison with the walking of humans, including infants.     

Methane Content in the Bottom Sediments and Water Column of the Black Sea

The methane content in the bottom sediments and water column of the Black Sea was determined using various methods of desorption and analysis of gases and various methods of calculating their concentrations. The head-space method with the use of salting out and calculation by an internal standard proved to be the most accurate procedure for the analysis of methane concentration in bottom sediments. The methane content in bottom sediments increased downward along the sediment thickness. In the upper 50–70 cm of shelf sediments, two minimums of methane concentration were revealed; in deep-sea sediments, only one minimum was recorded (in the 20–50 cm horizons). In the water column, methane concentrations slowly grew from the surface to a depth of 150–200 m and abruptly increased to a depth of 700–1200 m, remaining virtually constant in underlying layers. In certain deep-sea regions, peaks of methane content in the 1000–1200 m horizons of the water column were revealed, which were most probably due to local influx of abyssal waters enriched with this gas.     

Intermittent Swimming by Mammals: A Strategy for Increasing Energetic Efficiency During Diving

The evolutionary history of marine mammals involved marked physiologicaland morphological modifications to change from terrestrial toaquatic locomotion. A consequence of this ancestry is that swimmingis energetically expensive for mammals in comparison to fish.This study examined the use of behavioral strategies by marinemammals to circumvent these elevated locomotor costs duringhorizontal swimming and vertical diving. Intermittent formsof locomotion, including wave-riding and porpoising when nearthe water surface, and prolonged gliding and a stroke and glidemode of propulsion when diving, enabled marine mammals to increasethe efficiency of aquatic locomotion. Video instrumentationpacks (8-mm camera, video recorder and time-depth microprocessor)deployed on deep diving bottlenose dolphins (Tursiops truncatus),northern elephant seals (Mirounga angustirostris), and Weddellseals (Leptonychotes weddellii) revealed exceptionally longperiods of gliding during descent to depth. Glide duration dependedon depth and represented nearly 80% of the descent for divesexceeding 200 m. Transitions in locomotor mode during divingwere attributed to buoyancy changes with compression of thelungs at depth, and were associated with a 9–60% reductionin the energetic cost of dives for the species examined. Bychanging to intermittent locomotor patterns, marine mammalsare able to increase travelling speed for little additionalenergetic cost when surface swimming, and to extend the durationof submergence despite limitations in oxygen stores when diving.     

Damaging UV radiation and invertebrate predation: conflicting selective pressures for zooplankton vertical distribution in the water column of low DOC lakes

In nature most organisms have to manage conflicting demands of food gathering, predator avoidance, and finding a favorable abiotic environment (oxygen, temperature, etc.) in order to maximize their fitness. In the vertical water column of lakes with high solar ultraviolet radiation (UV) and invertebrate predators, zooplankton face two particularly strong and conflicting selective pressures. During daylight hours invertebrate predators often induce an upward vertical migration of zooplankton prey while potentially damaging UV forces a downward migration. We used 2.2 m long columns suspended vertically in a lake to conduct 2×2 factorial experiments to examine patterns of depth selection behavior by zooplankton in the presence and absence of both the invertebrate predator Chaoborus and UV. We hypothesized that Chaoborus and UV both affect the distribution of zooplankton and a combination of both factors would lead to a narrowing of depth distribution. We found that when Chaoborus were present zooplankton tended to be distributed at shallower depths in the columns, while in the presence of UV they exhibited a deeper distribution. Chaoborus themselves were always found near the bottom of the columns regardless of the UV treatment. Simultaneous exposure to predators and UV resulted in a peak of zooplankton (especially Daphnia catawba) distribution at intermediate depths. In a significant number of cases, depth range was narrowed in response to Chaoborus, UV, or both.     

Comparative photobehavior of marine cercariae with differing secondary host preferences

Light may serve as an important exogenous cue for parasitic larvae that have multi-host lifecycles and need to locate specific microhabitats, thereby increasing the probability of encountering their next host. We studied light as an initiating and orienting cue for swimming in two species of marine cercariae (Trematoda), Euhaplorchis sp. and Probolocoryphe lanceolata, which initially parasitize the same species of benthic snail, but then utilize different second intermediate hosts located in pelagic and benthic habitats, respectively. When tested in a laboratory simulation of underwater angular light distribution, dark-adapted Euhaplorchis cercariae swam slowly in darkness but ascended quickly toward downwelling light at quantal intensities over 4.0 × 10(15) photons m(-2) s(-1). They oriented toward a directional light source in a horizontal trough, confirming that light plays both an initiating and an orienting role in phototactic behavior that results in ascent in the water column to locate a fish host. In contrast, Probolocoryphe lanceolata cercariae exhibited haphazard vertical swimming in darkness, with downward swimming upon exposure to angular light at lower quantal intensities (>4.0 × 10(14) photons m(-2) s(-1)) than initiated swimming in Euhaplorchis. However, P. lanceolata cercariae did not swim in response to a directional light source, suggesting that while light initiated descent behavior, its orientation was controlled by another factor. These differences in photobehavior support the idea that trematode cercariae use light in selecting for microhabitats frequented by potential hosts: an adaptive benefit that optimizes their contact and transmission to the next host.     

Diving patterns and performance in the Antarctic blue-eyed shag Phalacrocorax atriceps

The pattern and characteristics of diving of two male blue-eyed shags Phalacrocorax atriceps were studied, using continuous-recording time-depth recorders, for a total of 15 consecutive days during which the depth, duration, bottom time, ascent and descent rates and surface intervals of 674 dives were recorded. Deep dives (> 35 m, averages80–90 m, max. 116 m) were twice as common (64% versus 34%) as shallow dives (< 21 m and 90% < 10 m). Deep dives were long (averages 2.7-4.1 min, max. 5.2 min) with half the time spent near maximum depth and fast travel speeds (averages 1.0-2.4 m s⁻¹). Shallow dives were short (average 0.5 min, max. 1.3 min), without bottom time and with slow travel speeds (0.1–0.6 m s⁻¹). The time spent at depth and the diet (mainly benthic fish and octopus) is consistent with benthic foraging; the function of shallow dives is uncertain. Male shags forage mainly in the afternoon in3–5 distinct bouts of diving. Within bouts (and shorter homogeneous sequences of diving) surface intervals are consistently2–3 times the preceding dive duration; in other shags the reverse is the case. Blue-eyed shag diving depth, duration and pattern is extreme amongst shags; and the relationship between dives and surface intervals suggests that they may regularly exceed their aerobic dive limit.