Dive Performance and Aquatic Thermoregulation of the World’s Smallest Mammalian Diver, the American Water Shrew (Sorex palustris)

AbstractAllometry predicts that the 12–17 g American water shrew (Sorex palustris)—the world’s smallest mammalian diver—will have the highest diving metabolic rate (DMR) coupled with the lowest total body oxygen storage capacity, skeletal muscle buffering capacity, and glycolytic potential of any endothermic diver. Despite these constraints, the maximum dive time (23.7 sec) and calculated aerobic dive limit (cADL; 10.8–14.4 sec) of wild-caught water shrews match or exceed values predicted by allometry based on studies of larger-bodied divers. The mean voluntary dive time of water shrews in 3, 10, 20, and 30°C water was 5.1±0.1 sec (N=25, n=1584), with a mean maximum dive time of 10.3±0.4 sec. Only 2.3–3.9% of dives in 30 and 10°C water, respectively, exceeded the cADL. Mean dive duration, duration of the longest dive, and total time in water all decreased significantly as temperature declined, suggesting that shrews employed behavioural thermoregulation to defend against immersion hypothermia. As expected from their low thermal inertia, diving shrews had a significantly higher DMR in 10°C (8.77 mL O2 g-1 hr-1) compared to 30°C water (6.57 mL O2 g-1 hr-1). Diving behavior of radio-telemetered shrews exclusively foraging in a simulated riparian environment (3°C water) for 12- to 28-hours suggest that mean (but not maximum) dive times of water shrews in the wild may be longer than predicted from our voluntary dive trials, as the average dive duration (6.9±0.2 sec, n=257) was ∼1.75× longer than during 20-min trials with no access to food at the same water temperature. Notably, free-diving shrews in the 24-hr trials consistently elevated core body temperature by ∼1.0–1.5°C immediately prior to initiating aquatic foraging bouts, and ended these bouts when body temperature was still at or above normal resting levels (∼37.8°C). We suggest this observed pre-dive hyperthermia aids to heighten the impressive somatosensory physiology, and hence foraging efficiency, of this diminutive predator while submerged.

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The development of diving behavior in juvenile Weddell seals: pushing physiological limits in order to survive

Canadian Journal of Zoology ◽

10.1139/z99-022 ◽

1999 ◽

Vol 77 (5) ◽

pp. 737-747 ◽

Cited By ~ 71

Author(s):

Jennifer M Burns

Keyword(s):

Blood Chemistry ◽

Weddell Seal ◽

Dive Duration ◽

Foraging Strategies ◽

Foraging Efficiency ◽

Diving Behavior ◽

Weddell Seals ◽

Physiological Limits ◽

Behavioral Constraints ◽

Absolute Energy

In juvenile phocids, the successful transition from nursing to independent foraging is contingent upon the development of adequate diving skills within the limited time between weaning and the depletion of body reserves. Yet, because juvenile seals are unable to remain submerged for as long as adults, owing to their smaller size, higher metabolic rates, and lowered oxygen stores, their behavioral options are likely constrained. To determine how such limitations might influence foraging strategies, we studied the development of diving behavior and physiology in Weddell seal (Leptonychotes weddellii) juveniles, using a combination of time-depth recorders, satellite-linked dive recorders, and morphological and physiological measurements (mass and blood chemistry). Time-depth recorder data indicated that the average depth, duration, and frequency of dives made by pups increased rapidly in the period from birth through weaning, but slowed soon thereafter. While preweaning increases in these parameters were correlated with seal age, postweaning increases in dive capacity were gradual and were probably the result of slower changes in mass and body composition. In weaned pups and yearlings, dive frequency and time underwater increased with age and (or) mass. Despite their smaller size and lower absolute energy requirements, the amount of time juveniles spent in the water was similar to that spent by adults. However, because juveniles were unable to remain submerged as long as adults and because most foraging dives were deep, juveniles were unable to spend an equivalent amount of time at the foraging depths. This difference was evident even though juveniles dove much closer to their anaerobic threshold than did adults. These findings support the hypothesis that the foraging efficiency of younger seals is reduced relative to that of adults, owing to physiological and morphological constraints on aerobic dive duration, and suggests that low juvenile survival might result from behavioral constraints.

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Body oxygen stores, aerobic dive limits and diving behaviour of the star-nosed mole (Condylura cristata) and comparisons with non-aquatic talpids

Journal of Experimental Biology ◽

10.1242/jeb.205.1.45 ◽

2002 ◽

Vol 205 (1) ◽

pp. 45-54

Author(s):

Ian W. McIntyre ◽

Kevin L. Campbell ◽

Robert A. MacArthur

Keyword(s):

Skeletal Muscles ◽

Storage Capacity ◽

Metabolic Cost ◽

Oxygen Storage Capacity ◽

Dive Duration ◽

Oxygen Storage ◽

High Mass ◽

The Mean ◽

Total Body ◽

Condylura Cristata

SUMMARY The dive performance, oxygen storage capacity and partitioning of body oxygen reserves of one of the world’s smallest mammalian divers, the star-nosed mole Condylura cristata, were investigated. On the basis of 722 voluntary dives recorded from 18 captive star-nosed moles, the mean dive duration (9.2±0.2 s; mean ± s.e.m.) and maximum recorded dive time (47 s) of this insectivore were comparable with those of several substantially larger semi-aquatic endotherms. Total body O2 stores of adult star-nosed moles (34.0 ml kg–1) were 16.4 % higher than for similarly sized, strictly fossorial coast moles Scapanus orarius (29.2 ml kg–1), with the greatest differences observed in lung and muscle O2 storage capacity. The mean lung volume of C. cristata (8.09 ml 100 g–1) was 1.81 times the predicted allometric value and exceeded that of coast moles by 65.4 % (P=0.0001). The overall mean myoglobin (Mb) concentration of skeletal muscles of adult star-nosed moles (13.57±0.40 mg g–1 wet tissue, N=7) was 19.5 % higher than for coast moles (11.36±0.34 mg g–1 wet tissue, N=10; P=0.0008) and 54.2 % higher than for American shrew-moles Neurotrichus gibbsii (8.8 mg g–1 wet tissue; N=2). The mean skeletal muscle Mb content of adult star-nosed moles was 91.1 % higher than for juveniles of this species (P<0.0001). On the basis of an average diving metabolic rate of 5.38±0.35 ml O2 g–1 h–1 (N=11), the calculated aerobic dive limit (ADL) of star-nosed moles was 22.8 s for adults and 20.7 s for juveniles. Only 2.9 % of voluntary dives by adult and juvenile star-nosed moles exceeded their respective calculated ADLs, suggesting that star-nosed moles rarely exploit anaerobic metabolism while diving, a conclusion supported by the low buffering capacity of their skeletal muscles. We suggest that a high mass-specific O2 storage capacity and relatively low metabolic cost of submergence are key contributors to the impressive dive performance of these diminutive insectivores.

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Blood viscosity and optimal hematocrit in a deep-diving mammal, the northern elephant seal (Mirounga angustirostris)

Canadian Journal of Zoology ◽

10.1139/z86-317 ◽

1986 ◽

Vol 64 (10) ◽

pp. 2081-2085 ◽

Cited By ~ 31

Author(s):

Michael S. Hedrick ◽

Deborah A. Duffield ◽

Lanny H. Cornell

Keyword(s):

Oxygen Transport ◽

Blood Viscosity ◽

Storage Capacity ◽

Oxygen Storage Capacity ◽

Elephant Seal ◽

Oxygen Storage ◽

Diving Behavior ◽

Northern Elephant Seal ◽

Elephant Seals ◽

Deep Diving

Elephant seals offer a unique opportunity to examine rheological characteristics of blood because of the normally high hematocrits in this species. A comparison of blood viscosity of the elephant seal with that of a terrestrial mammal (rabbit; HCT = 35%) reveals a threefold increase in viscosity of elephant seal blood over that of rabbit blood due to the high hematocrit (HCT = 65%). While the increased hematocrit of elephant seal blood reflects increased oxygen storage capacity, blood oxygen transport may actually be reduced by the effects of increased blood viscosity on blood flow. Elephant seal plasma viscosity was also higher than that of rabbit plasma; this was associated with a higher concentration of plasma proteins. There were no apparent differences in the viscous properties of the red blood cells of the two species. The theoretically optimal hematocrit was determined in vitro for reconstituted blood from each species and compared with the observed in vivo hematocrit. It was found that the observed hematocrit of the elephant seal lies far to the right of the predicted hematocrit for optimal oxygen transport, while the rabbit hematocrit was identical with the predicted value. These results suggest that elephant seals have increased oxygen storage capacity at the expense of optimizing oxygen transport. The observed increase in hematocrit and viscosity may be of importance in considering the diving behavior and energetics of elephant seals.

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Diving performance of male and female Japanese Cormorants

Canadian Journal of Zoology ◽

10.1139/z96-122 ◽

1996 ◽

Vol 74 (6) ◽

pp. 1098-1109 ◽

Cited By ~ 47

Author(s):

Yutaka Watanuki ◽

Akiko Kato ◽

Yasuhiko Naito

Keyword(s):

Body Mass ◽

Dive Duration ◽

Sexual Differences ◽

Foraging Efficiency ◽

Diving Behavior ◽

Bottom Phase ◽

Similar Proportion ◽

Sexually Dimorphic ◽

Surface Time ◽

Ascent Phase

Sexual differences in the diving behavior of the sexually dimorphic Japanese Cormorant, Phalacrocorax capillatus (males are 26% heavier than females), were studied at Teuri Island, Hokkaido, using time–depth recorders. A typical dive cycle involved a rapid descent phase, a bottom phase where they remained for a while, an ascent phase, and a postdive surface phase. Depth and duration across individual birds were greater for males (15.1 ± 3.7 (mean ± SD) m, 37 ± 5 s, respectively) than those for females (7.2 ± 2.4 m, 24 ± 4 s, respectively). While submerged, females spent a similar proportion of time during the bottom phase to males, hence foraging efficiency (proportion of time at the bottom to total dive cycle time) did not differ between the sexes. No sexual differences were found in descent and ascent rates, dive bout duration, or time spent underwater per day. No significant effects of dive duration on postdive surface time were observed for either sex, indicating that birds dived within an aerobic dive limit. However, mean dive durations and maximum dive durations for individual birds were a function of body mass to the power 1.49 and 1.87, respectively, suggesting that body mass partly constrains the diving behavior of this opportunistically feeding cormorant.

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Gas exchange of captive freely diving grey seals (Halichoerus grypus).

Journal of Experimental Biology ◽

10.1242/jeb.191.1.1 ◽

1994 ◽

Vol 191 (1) ◽

pp. 1-18 ◽

Cited By ~ 7

Author(s):

J Z Reed ◽

C Chambers ◽

M A Fedak ◽

P J Butler

Keyword(s):

Gas Exchange ◽

Oxygen Storage Capacity ◽

Dive Duration ◽

Minute Volume ◽

Oxygen Storage ◽

Allometric Relationships ◽

Aerobic Dive Limit ◽

Oxygen Utilisation ◽

End Tidal ◽

Limited Oxygen

When at sea, phocids dive for long periods and spend a high percentage of their time submerged. This behaviour requires some combination of an increased oxygen storage capacity, rapid oxygen loading at the surface and reduced oxygen utilisation when submerged. To assess these adaptations, breath-by-breath ventilation was studied in four adult grey seals (two male, two female, 160-250 kg), freely diving in a large outdoor tank where surface access was restricted to one breathing hole. The dive patterns obtained were similar to those recorded from freely diving wild grey seals. Respiratory frequency during the surface periods was 40% higher than that estimated from allometric relationships (19.4 +/- 0.7 breaths min-1), and tidal volume (6.3 +/- 1.21) was approximately five times higher than that estimated from allometric relationships. These adaptations produce a high minute volume and enable gas exchange to occur at the surface. Mean oxygen consumption rate (VO2, measured for a dive+surface cycle) decreased with increasing dive duration. The aerobic dive limit was estimated as 9.6 min for a 150 kg grey seal (using the overall average VO2 of 5.2 ml O2 min-1 kg-1), which is consistent with results from freely diving wild grey seals (only 6% of dives exceeded 10 min). End-tidal oxygen values varied during a surface period, following a U-shaped curve, which suggests that there is limited oxygen uptake from the lung and/or blood oxygen stores during dives. This result was unexpected and indicates that these seals are utilising substantial physiological responses to conserve oxygen, even during shallow voluntary diving.

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