HEART RATE AND ELECTROCARDIOGRAM CHARACTERISTICS OF A YOUNG CALIFORNIA GRAY WHALE (ESCHRICHTIUS ROBUSTUS)1

Electrocardiogram (ECG) analyses of Holter monitor recordings from a young California gray whale were performed to determine ECG waveform characteristics, evaluate the heart rate pattern for sinus arrhythmia, obtain resting heart rates at known body masses as the whale increased in size, and compare those heart rates with predicted heart rates from allometric equations. The PR and QRS intervals (475 ± 35 msec, 208 ± 24 msec, respectively, n= 20) support the concept (Meijler et al. 1992) that atrioventricular transmission and ventricular excitation times do not increase linearly in very large mammals. A sinus arrhythmia pattern at rest (apneic heart rates of 15–25 beats per min [bpm] and eupneic heart rates of 34–40 bpm) is consistent with a relative eupneic tachycardia and apneic bradycardia during diving activity of whales. The heart rate-body mass measurements (35–24 bpm at body masses of 3,591-8,200 kg) in this study (1) extend the range of allometric heart rate and body mass data in mammals a full order of magnitude, to almost 10,000 kg, (2) support the use of allometric equations (based primarily on mammals <1,000 kg in body mass) in estimating resting heart rates in whales, and (3) demonstrate that previously reported heart rates in large whales are not representative of resting heart rate, probably secondary to circumstances during measurement.     

Measurement of the CO2 apneic threshold in newborn infants: possible relevance for periodic breathing and apnea.

We measured the PCO2 apneic threshold in preterm and term infants. We hypothesized that, compared with adult subjects, the PCO2 apneic threshold in neonates is very close to the eupneic PCO2, likely facilitating the appearance of periodic breathing and apnea. In contrast with adults, who need to be artificially hyperventilated to switch from regular to periodic breathing, neonates do this spontaneously. We therefore measured the apneic threshold as the average alveolar PCO2 (PaCO2) of the last three breaths of regular breathing preceding the first apnea of an epoch of periodic breathing. We also measured the PaCO2 of the first three breaths of regular breathing after the last apnea of the same periodic breathing epoch. In preterm infants, eupneic PaCO2 was 38.6 +/- 1.4 Torr, the preperiodic PaCO2 apneic threshold was 37.3 +/- 1.4 Torr, and the postperiodic PaCO2 was 37.2 +/- 1.4 Torr. In term infants, the eupneic PaCO2 was 39.7 +/- 1.1 Torr, the preperiodic PaCO2 apneic threshold was 38.7 +/- 1.0 Torr, and the postperiodic value was 37.9 +/- 1.2 Torr. This means that the PaCO2 apneic thresholds were 1.3 +/- 0.1 and 1.0 +/- 0.2 Torr below eupneic PaCO2 in preterm and term infants, respectively. The transition from eupneic PaCO2 to PaCO2 apneic threshold preceding periodic breathing was accompanied by a minor and nonsignificant increase in ventilation, primarily related to a slight increase in frequency. The findings suggest that neonates breathe very close to their PCO2 apneic threshold, the overall average eupneic PCO2 being only 1.15 +/- 0.2 Torr (0.95-1.79, 95% confidence interval) above the apneic threshold. This value is much lower than that reported for adult subjects (3.5 +/- 0.4 Torr). We speculate that this closeness of eupneic and apneic PCO2 thresholds confers great vulnerability to the respiratory control system in neonates, because minor oscillations in breathing may bring eupneic PCO2 below threshold, causing apnea.     

DEVELOPMENT OF A BIMODAL HEART RATE PATTERN IN UNDISTURBED FETAL HARBOR SEALS

Fetal and maternal heart rates were studied in unrestrained, pregnant harbor seals during the last third of gestation. Heart rates were recorded while the mothers were resting on land or performing trained simulated dives of up to 2.25 min. Data from resting mothers showed the development of a bimodal or two-speed fetal heart rate pattern during late gestation. The mean faster and slower fetal heart rates at term were 125 ± 3.7 and 79 ± 3.1 (mean + SEM) respectively. The amount of fetal bradycardia observed increased steadily towards term, and fetal heart rate changes were not correlated with changes in maternal heart rate or maternal respiration. The bimodal fetal heart rate was also seen during the simulated dives, and no decrease in either the faster or slower heart rate was found. Heart rates from resting, unrestrained harbor seal pups were also studied. The pups displayed a bimodal heart rate similar to the fetuses' with the slower rate occurring during breath-holds. The bradycardia in the pups was equivalent to the slower fetal heart rate. These findings suggest that the regulatory mechanism that determines the apneic bradycardia in young harbor seals during non-stressful conditions develops in the last quarter of gestation.     

Cardiac output and muscle blood flow during rest-associated apneas of elephant seals

In order to evaluate hemodynamics and blood flow during rest-associated apnea in young elephant seals (Mirounga angustirostris), cardiac outputs (CO, thermodilution), heart rates (HR), and muscle blood flow (MBF, laser Doppler flowmetry) were measured. Mean apneic COs and HRs of three seals were 46% and 39% less than eupneic values, respectively (2.1+/-0.3 vs. 4.0+/-0.1 mL kg(-1) s(-1), and 54+/-6 vs. 89+/-14 beats min(-1)). The mean apneic stroke volume (SV) was not significantly different from the eupneic value (2.3+/-0.2 vs. 2.7+/-0.5 mL kg(-1)). Mean apneic MBF of three seals was 51% of the eupneic value. The decline in MBF during apnea was gradual, and variable in both rate and magnitude. In contrast to values previously documented in seals during forced submersions (FS), CO and SV during rest-associated apneas were maintained at levels characteristic of previously published values in similarly-sized terrestrial mammals at rest. Apneic COs of such magnitude and incomplete muscle ischemia during the apnea suggest that (1) most organs are not ischemic during rest-associated apneas, (2) the blood O(2) depletion rate is greater during rest-associated apneas than during FS, and (3) the blood O(2) store is not completely isolated from muscle during rest-associated apneas.     

Movements and diving behavior of weaned Weddell seal (Leptonychotes weddellii ) pups

Between 1993 and 1995, the diving behavior and movement patterns of 23 weaned Weddell seal pups (Leptonychotes weddellii) were tracked in the Ross Sea. Antarctica, using satellite-linked time-depth recorders. Regression analyses revealed that for seals of between 8 and 27 weeks old, age was poorly correlated with the dive depth, duration, or frequency. However, changes in dive parameters suggested that Weddell seal pups were attempting to maximize dive time, but the manner in which this was done depended on age and time of day. Movement patterns indicated that most Weddell seal pups left their natal area by the end of February, and traveled north along the Antarctic continent coastline. Several individuals returned to McMurdo Sound, but others were last located more than 400 km from McMurdo. Routes followed suggest that pups can use the pack ice habitat, but prefer to remain closer to the coastline than do adults. Accepted: 21 July 1998     

Weddell seals do not lengthen calls in response to conspecific masking

Some mammalian and avian species alter their vocal communication signals to reduce masking by background noises (including conspecific calls). A preliminary study suggested that Weddell seals (Leptonychotes weddellii) increase the durations of some underwater call types when overlapped by another calling seal. The present study examined the durations and overlapping sequences of Weddell seal calls recorded in Eastern Antarctica. The calling rate, call type (13 major categories), total duration, numbers of elements per call and overlapping order of 100–200 consecutive calls per recording location were measured. In response to increased conspecific calling rates, the call durations and numbers of elements (within repeated-element call types) did not change or became shorter. Calls that were not overlapped were 3.8?±?6.1 s long, the first call in a series of overlapped calls was 14.4?±?15.7 s and subsequent calls in an overlapping series were 6.5?±?10.3 s. The mean durations of non-overlapped and overlapped calls matched random distributions. Weddell seals do not appear to be adjusting the durations or timing of their calls to purposefully avoid masking each other’s calls. The longer a call is, the more likely it is to overlap another call by chance. An implication of this is that Weddell seals may not have the behavioural flexibility to reduce masking by altering the temporal aspects of their calls or calling behaviours as background noises (natural and from shipping) increase.     

Underwater calling rates of harp and Weddell seals as a function of hydrophone location

Stereo recordings of harp seal (Pagophilus groenlandicus) and Weddell seal (Leptonychotes weddellii) underwater vocalizations were made near breeding groups using separations between two hydrophones of 159–339 m. Within a large herd, harp seal call numbers varied slightly between channels. Counts of Weddell seal calls were higher near a small herd on the ice than 159–180 m away. Repeat counts of harp seal calls by a single observer differed significantly. Source levels of Weddell seal calls varied and higher amplitude calls would be detected up to 2 orders of magnitude farther away than quieter calls. Hydrophone location (especially near small groups), observer variability and call source level differences will bias the use of monitoring underwater seal vocalizations to index locations, population size or underwater behaviours. Using hydrophone arrays and multiple observers may mitigate these problems. Accepted: 15 September 2000     

Effects of increase in body temperature on the breathing pattern in premature infants

This study was designed to determine the effects of a mild increase in body temperature within the physiological range (0.8 degrees C) in healthy premature infants. Seven unsedated premature infants (38.4 wk +/- 1.5 postconceptional age) were monitored polygraphically during "morning naps" in an incubator under two different environmental temperatures: (1) normothermia with the incubator temperature set at 25 degrees C and the rectal temperature equal to 36.9 degrees C +/- 0.1; (2) hyperthermia with the incubator temperature set at 35 degrees C and the rectal temperature equal to 37.7 degrees C +/- 0.15. Respiratory frequency and heart rate, respiratory events, i.e., central and obstructive apnea, and periodic breathing with and without apneic oscillations were tabulated. Results for respiratory events were expressed as (1) indices of the total number of respiratory events, and of specific respiratory events per hour of total, quiet and active sleep times; (2) duration of total and specific respiratory events expressed as a percentage of total sleep, quiet and active sleep times. Respiratory frequency and heart rate were significantly increased by hyperthermia (P less than 0.05). Hyperthermia did not significantly modify the indices or the duration of central and obstructive apnea. But the indices and the duration of periodic breathing with and without apneic oscillations were significantly increased by hyperthermia during active sleep (P less than 0.05) but not during quiet sleep. The present study shows that a mild increase in body temperature within the physiological range in premature infants enhances the instability of the breathing pattern during active sleep.     

PHYSIOLOGY WITHOUT RESTRAINT IN DIVING MAMMALS

Forced submersions and natural dives are compared with regard to heart rate, blood distribution and O₂ consumption. Heart rate has been used as a primary indicator of blood distribution. However, often the dive heart rate has been compared to inappropriate surface conditions, and it is suggested that it should be compared to resting apneusis in the arrhythmic breathing animal. Furthermore, natural dives are frequently short relative to the breathhold capacity of a diver, and often during these dives there is little change in heart rate or apparent kidney function from the resting state. Calculations are made to determine the breathhold limit without O₂ conservation measures. Further calculations are made to estimate the expected metabolism and metabolite production during an exceptionally long dive in a seal when O₂ conservation must be invoked. These results are compared to known values obtained from a Weddell seal that made a 61 min dive. Based on the results of these observations and calculations, three possible models of blood distribution during natural dives are discussed. This discussion shows the limitation of present data and the type of data needed to resolve the question of how aquatic vertebrates manage O₂ stores during dives of varying durations.     

Some anatomical aspects of the cardiovascular system of antarctic seals and their possible functional significance in diving

The hearts and ascending aortae of 11 Weddell seals, Leptonychotes weddelli, three adult Crabeater seals, Lobodon carcinophagus, two adult Ross seals, Ommatophoca rossi, and one adult Leopard seal, Hydrurga leptonyx, were examined for comparison with terrestrial forms. The Weddell seal specimens were from animals ranging in age from midterm in fetal development to mature adults. All specimens were collected in 1971, 1972, and 1973, from McMurdo Sound and the Ross Sea, Antarctica. The phocid hearts were characteristically broader and flatter than those of other carnivore families and they tended toward bifid apices. The heart form indices (height/circumference) averaged 31.5 compared to 39.0 for felids. The right ventricular chambers of the Antarctic seals were found to average longer in Weddells and narrower in all, than those reported for four other carnivore families. An elastic enlargement was present in the ascending aortae of all seals. The largest diameter of the aortic bulb averaged 25.5 mm more than the base of the aorta in the adult Weddell seals which represented an increase of 72.5% over the base. It is suggested that the general heart form, and especially the ascending aortae, are anatomical adaptations to diving. The compressed heart makes possible unimpaired function when the chest is compressed during deep dives. The aortic bulb maintains mean arterial blood pressure and perfusion of the brain and cardiac tissue during diving bradycardia.     

Transformation of metabolism with age and lifestyle in Antarctic seals: a case study of systems biology approach to cross-species microarray experiment

Background  

The metabolic transformation that changes Weddell seal pups born on land into aquatic animals is not only interesting for the study of general biology, but it also provides a model for the acquired and congenital muscle disorders which are associated with oxygen metabolism in skeletal muscle. However, the analysis of gene expression in seals is hampered by the lack of specific microarrays and the very limited annotation of known Weddell seal (Leptonychotes weddellii) genes.     

Histological,histochemical, and ultrastructural investigations on the gastrointestinal system of antarctic seals: Weddell seal (Leptonychotes weddellii) and crabeater seal (Lobodon carcinophagus)

The morphology of the principal sections of the gastrointestinal system of two Antarctic seals with different dietary habits, namely, the Weddell seal (Leptonychotes weddellii) and the crabeater seal (Lobodon carcinophagus), has been investigated. Histologically examined by light microscopy, the tissue layers of the gastrointestinal tract of both seals are almost identical to those observed in most other mammals and no major differences in principle organization could be found between the two seal species. The ultrastructure of the gastric and intestinal epithelial cells has been examined and is also closely comparable to that of these cells in other mammals; however, Paneth cells have not been found in our material. In general, therefore, adaptations of the gastrointestinal tract to the aquatic environment or the diet are not obvious at the morphological levels of organization studied. Histochemical differences are found between the two closely related species; mucins of the surface epithelium in the stomach of Weddell seals are highly sulfated, while those in the crabeater seal are not. Mucous neck cells in Weddell seals contain acid mucosubstances, while those of crabeater seals contain neutral ones. Goblet cells in the small and large intestine in Weddell seals contain both neutral and acid mucosubstances. Both mucin types are detected in the crabeater seal; however, the mucins of the colon in the crabeater seal are more highly sulfated than those in the Weddell seal. The ratio of globet cells to enterocytes in the large intestine of crabeater seals is higher than that in Weddell seals. © 1995 Wiley-Liss, Inc.     

Feeding habits of three seal species at the Danco Coast, Antarctica: a re-assessment

The analysis of prey overlap among Weddell, Antarctic fur and leopard seals was conducted using fecal samples collected at the Danco Coast, Antarctic Peninsula, in 1998 and 2000. The re-occurrence of prey species was moderate in samples collected in 1998, and low in 2000, and reflects resource partitioning among seal species. Prey species that mostly co-occurred in seals’ diet were the Antarctic krill Euphausia superba, bivalves, and the myctophids Gymnoscopelus nicholsi and Electrona antarctica. A dietary similarity index of prey overlap has been calculated and demonstrates evident fluctuations in pairwise comparisons between the seal species. The highest and lowest values of prey overlap were observed between Antarctic fur seals and leopard seals, and between Weddell seals and leopard seals, respectively. Prey overlap between Antarctic fur seals and Weddell seals was moderate in both seasons.     

Dinucleotide microsatellite markers from the Antarctic seals and their use in other Pinnipeds

Twenty‐four microsatellite loci were isolated from three species of Antarctic seals (Subfamily Monachinae, Tribe Lobodontini). Eleven loci were cloned from Weddell seal, Leptonychotes weddellii, seven from leopard seal, Hydrurga leptonyx, and six from crabeater seal, Lobodon carcinophagus. Variability was assessed in Weddell seals collected in McMurdo Sound, leopard seals from Bird Island, South Georgia, and crabeater seals sampled in the eastern Ross Sea. All loci were variable in the three species used for cloning and 22 of these loci amplified variable products in the Ross seal, Ommatophoca rossii. Cross‐species amplification was largely successful, with an average of 19 loci amplifying products in other phocids.     

Contrasting behavior between two populations of an ice‐obligate predator in East Antarctica

The Austral autumn–winter is a critical period for capital breeders such as Weddell seals that must optimize resource acquisition and storage to provision breeding in the subsequent spring. However, how Weddell seals find food in the winter months remains poorly documented. We equipped adult Weddell seals after their annual molt with satellite‐relayed data loggers at two sites in East Antarctica: Dumont D'Urville (n = 12, DDU) and Davis (n = 20). We used binomial generalized mixed‐effect models to investigate Weddell seals’ behavioral response (i.e., “hunting” vs. “transit”) to physical aspects of their environment (e.g., ice concentration). Weddell seal foraging was concentrated to within 5 km of a breathing hole, and they appear to move between holes as local food is depleted. There were regional differences in behavior so that seals at Davis traveled greater distances (three times more) and spent less time in hunting mode (half the time) than seals at DDU. Despite these differences, hunting dives at both locations were pelagic, concentrated in areas of high ice concentration, and over areas of complex bathymetry. There was also a seasonal change in diving behavior from transiting early in the season to more hunting during winter. Our observations suggest that Weddell seal foraging behavior is plastic and that they respond behaviorally to changes in their environment to maximize food acquisition and storage. Such plasticity is a hallmark of animals that live in very dynamic environments such as the high Antarctic where resources are unpredictable.     

Geographic variation of Weddell seal (<Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>) airborne mother–pup vocalisations

Geographic variation of vocal behaviour may be reflective of genetic, cultural and ecological differences between locations. Such differences have been previously reported in six pinnipeds, including Weddell seals (Leptonychotes weddellii). This study investigated whether the acoustic characteristics of airborne Weddell seal mother and pup ‘contact’ calls vary geographically between the Vestfold Hills and McMurdo Sound. From each location, 72 mother and 120 pup calls were analysed, with 11 acoustic characteristics measured. MANOVA and Discriminant function analysis (DFA) identified significant differences in several fundamental frequency characteristics. The cross-validated DFA classified greater than 60% of mother and pup vocalisations to the correct location. For novel calls, the probabilities of correctly classifying these percentages by chance are low. Additionally, ordinal logistic regression identified significant differences in the maximum energy distribution of mother and pup vocalisations. For pups, this variation is likely the result of genetic differences between the two populations. Nevertheless, results support suggestions that, when compared with underwater repertoire differences, airborne within-call differences are less efficient at identifying Weddell seal breeding populations.     

Summer diet of Weddell Seals (Leptonychotes weddelli) in the eastern and southern Weddell Sea,Antarctica

Summary Stomach and intestine samples from 21 adult Weddell seals were used to study the diet of these seals from the eastern and southern Weddell Sea coast from January to February 1983 and 1985. Fish occurred in all seals, squid in five, octopods in three and Euphausia crystallorophias in one seal. Pleuragramma antarcticum was the predominant fish in the diet, constituting 61.1% of otoliths in 1983 samples and 93.8% in 1985. Aethotaxis mitopteryx, Dissostichus mawsoni, unidentified Trematomus spp. and channichthyids were also recorded. Size and wet weight of P. antarcticum were calculated from uneroded otoliths, found in 6 seal stomachs with liquid food pulp, collected during early morning hours in 1985. Size distribution of P. antarcticum from individual seals was reasonably constant, ranging between 5.0 and 22.0 cm SL; adult fish from about 14.0 to 19.0 cm SL predominated. P. antarcticum in seals from the southern area had a larger median size (16.5 cm SL), than those from further east (15.5 cm SL). Calculated wet weights of all P. antarcticum from individual seal stomachs ranged between 4.7 and 16.9 kg the mean was 12.8 kg. Comparisons with net-hauls from the southern Gould Bay suggest that Weddell seals feed mainly in deeper water layers (>400 m) where adult P. antarcticum occur at higher densities.     

Effect of gender on the development of hypocapnic apnea/hypopnea during NREM sleep.

We hypothesized that a decreased susceptibility to the development of hypocapnic central apnea during non-rapid eye movement (NREM) sleep in women compared with men could be an explanation for the gender difference in the sleep apnea/hypopnea syndrome. We studied eight men (age 25-35 yr) and eight women in the midluteal phase of the menstrual cycle (age 21-43 yr); we repeated studies in six women during the midfollicular phase. Hypocapnia was induced via nasal mechanical ventilation for 3 min, with respiratory frequency matched to eupneic frequency. Tidal volume (VT) was increased between 110 and 200% of eupneic control. Cessation of mechanical ventilation resulted in hypocapnic central apnea or hypopnea, depending on the magnitude of hypocapnia. Nadir minute ventilation in the recovery period was plotted against the change in end-tidal PCO(2) (PET(CO(2))) per trial; minute ventilation was given a value of 0 during central apnea. The apneic threshold was defined as the x-intercept of the linear regression line. In women, induction of a central apnea required an increase in VT to 155 +/- 29% (mean +/- SD) and a reduction of PET(CO(2)) by -4.72 +/- 0.57 Torr. In men, induction of a central apnea required an increase in VT to 142 +/- 13% and a reduction of PET(CO(2)) by -3.54 +/- 0.31 Torr (P = 0.002). There was no difference in the apneic threshold between the follicular and the luteal phase in women. Premenopausal women are less susceptible to hypocapnic disfacilitation during NREM sleep than men. This effect was not explained by progesterone. Preservation of ventilatory motor output during hypocapnia may explain the gender difference in sleep apnea.     

The effect of feeding on the metabolic rate in harbour seals (Phoca vitulina)

The heat increment of feeding was estimated in harbour seals (Phoca vitulina). Seals were given different amounts of herring, ranging from 0.8 to 2.65 kg. The caloric content of the herring ranged from 6575 to 12560 kJ·kg^-1 depending on time of year. Metabolic rate increased within 30 min after feeding, and the magnitude and duration of heat increment of feeding depended on the size of the meal and the caloric content of the herring. Measured heat increment of feeding was up to 14.9% of gross energy intake and metabolic rate increased as much as 46% above resting, postabsorptive metabolic rate for 15 h duration in a harbour seal with a body weight of approximately 40 kg.     

CLASSIFICATION OF DIVERSE CALL TYPES USING CLUSTER ANALYSIS TECHNIQUES

ABSTRACT

We investigated the problem of categorizing the repertoire of a group of highly varied vocalizations. A set of Weddell seal Leptonychotes weddellim-air calls recorded near Davis, Antarctica, were examined. The repertoire size was estimated by first subjectively assigning each call (based on auditory and spectrographic patterns) to one of a large number of provisional call types. A set of frequency, duration and waveform measurements were made on every call. For each provisional call type, the mean value of each of these measures was calculated. These mean values were used to perform Cluster Analyses of the provisional call types. Beginning with calls clustered closest together, the most similar provisional call types were successively amalgamated until further joining would link two clearly dissimilar vocalizations (rising versus falling frequency sweeps in this case). The Weddell seal repertoire contained 12 call types ranging from long sinusoidal upsweeps to growls. This procedure provides a method of estimating the minimum repertoire size of a sample of calls. Dichotomous Sorting using Principal Components Analysis can also be used to categorize calls but will probably be of greater value when applied to finding subdivisions within a single call type.