Non-auditory, electrophysiological potentials preceding dolphin biosonar click production.

The auditory brainstem response to a dolphin's own emitted biosonar click can be measured by averaging epochs of the instantaneous electroencephalogram (EEG) that are time-locked to the emitted click. In this study, averaged EEGs were measured using surface electrodes placed on the head in six different configurations while dolphins performed an echolocation task. Simultaneously, biosonar click emissions were measured using contact hydrophones on the melon and a hydrophone in the farfield. The averaged EEGs revealed an electrophysiological potential (the pre-auditory wave, PAW) that preceded the production of each biosonar click. The largest PAW amplitudes occurred with the non-inverting electrode just right of the midline-the apparent side of biosonar click generation-and posterior of the blowhole. Although the source of the PAW is unknown, the temporal and spatial properties rule out an auditory source. The PAW may be a neural or myogenic potential associated with click production; however, it is not known if muscles within the dolphin nasal system can be actuated at the high rates reported for dolphin click production, or if sufficiently coordinated and fast motor endplates of nasal muscles exist to produce a PAW detectable with surface electrodes.

History of the Development of Anesthesia for the Dolphin: A Quest to Study a Brain as Large as Man's.

It is important for academic-minded human anesthesiologists to have an interdisciplinary perspective when engaging in cutting-edge research as well as the practice of human anesthesiology. This was a philosophy promoted by Dr. Robert Dripps, former pioneering Chairman of the Anesthesiology Department at the University of Pennsylvania (Philadelphia, Pennsylvania). Many human and veterinary anesthesiologists as well as biomedical engineers and neuroscientists benefited from Dr. Dripps's constructive outlook personified in the quest to develop dolphin anesthesiology.The motivation to anesthetize dolphins came from the fact that scientists and physicians wanted to study the brain of the dolphin, a brain as large as man's. Also, investigators wanted to develop anesthesia for the dolphin in order to study the electrophysiology of the dolphin's highly sophisticated auditory system, which facilitates the dolphin's amazing echolocation capability.Dolphin anesthesia involves a complex matter of unique neural control, airway anatomy, neuromuscular control of respiration, and sleep behavior.

Comparison of Dolphins' Body and Brain Measurements with Four Other Groups of Cetaceans Reveals Great Diversity.

We compared mature dolphins with 4 other groupings of mature cetaceans. With a large data set, we found great brain diversity among 5 different taxonomic groupings. The dolphins in our data set ranged in body mass from about 40 to 6,750 kg and in brain mass from 0.4 to 9.3 kg. Dolphin body length ranged from 1.3 to 7.6 m. In our combined data set from the 4 other groups of cetaceans, body mass ranged from about 20 to 120,000 kg and brain mass from about 0.2 to 9.2 kg, while body length varied from 1.21 to 26.8 m. Not all cetaceans have large brains relative to their body size. A few dolphins near human body size have human-sized brains. On the other hand, the absolute brain mass of some other cetaceans is only one-sixth as large. We found that brain volume relative to body mass decreases from Delphinidae to a group of Phocoenidae and Monodontidae, to a group of other odontocetes, to Balaenopteroidea, and finally to Balaenidae. We also found the same general trend when we compared brain volume relative to body length, except that the Delphinidae and Phocoenidae-Monodontidae groups do not differ significantly. The Balaenidae have the smallest relative brain mass and the lowest cerebral cortex surface area. Brain parts also vary. Relative to body mass and to body length, dolphins also have the largest cerebellums. Cortex surface area is isometric with brain size when we exclude the Balaenidae. Our data show that the brains of Balaenidae are less convoluted than those of the other cetaceans measured. Large vascular networks inside the cranial vault may help to maintain brain temperature, and these nonbrain tissues increase in volume with body mass and with body length ranging from 8 to 65% of the endocranial volume. Because endocranial vascular networks and other adnexa, such as the tentorium cerebelli, vary so much in different species, brain size measures from endocasts of some extinct cetaceans may be overestimates. Our regression of body length on endocranial adnexa might be used for better estimates of brain volume from endocasts or from endocranial volume of living species or extinct cetaceans.

Non-Fermi-Liquid Behavior and Anomalous Suppression of Landau Damping in Layered Metals Close to Ferromagnetism.

We analyze the low-energy physics of nearly ferromagnetic metals in two spatial dimensions using the functional renormalization group technique. We find a new low-energy fixed point, at which the fermionic (electronlike) excitations are non-Fermi-liquid (z_{f}=13/10) and the magnetic fluctuations exhibit an anomalous Landau damping whose rate vanishes as Γ_{q}∼|q|^{3/5} in the low-|q| limit. We discuss this renormalization of the Landau-damping exponent, which is the major novel prediction of our work, and highlight the possible link between that renormalization and neutron-scattering data on UGe_{2} and related compounds. Implications of our analysis for YFe_{2}Al_{10} are also discussed.

On doing two things at once: dolphin brain and nose coordinate sonar clicks, buzzes and emotional squeals with social sounds during fish capture.

Dolphins fishing alone in open waters may whistle without interrupting their sonar clicks as they find and eat or reject fish. Our study is the first to match sound and video from the dolphin with sound and video from near the fish. During search and capture of fish, free-swimming dolphins carried cameras to record video and sound. A hydrophone in the far field near the fish also recorded sound. From these two perspectives, we studied the time course of dolphin sound production during fish capture. Our observations identify the instant of fish capture. There are three consistent acoustic phases: sonar clicks locate the fish; about 0.4 s before capture, the dolphin clicks become more rapid to form a second phase, the terminal buzz; at or just before capture, the buzz turns to an emotional squeal (the victory squeal), which may last 0.2 to 20 s after capture. The squeals are pulse bursts that vary in duration, peak frequency and amplitude. The victory squeal may be a reflection of emotion triggered by brain dopamine release. It may also affect prey to ease capture and/or it may be a way to communicate the presence of food to other dolphins. Dolphins also use whistles as communication or social sounds. Whistling during sonar clicking suggests that dolphins may be adept at doing two things at once. We know that dolphin brain hemispheres may sleep independently. Our results suggest that the two dolphin brain hemispheres may also act independently in communication.

Sperm whales and killer whales with the largest brains of all toothed whales show extreme differences in cerebellum.

Among cetaceans, killer whales and sperm whales have the widest distribution in the world's oceans. Both species use echolocation, are long-lived, and have the longest periods of gestation among whales. Sperm whales dive much deeper and much longer than killer whales. It has long been thought that sperm whales have the largest brains of all living things, but our brain mass evidence, from published sources and our own specimens, shows that big males of these two species share this distinction. Despite this, we also find that cerebellum size is very different between killer whales and sperm whales. The sperm whale cerebellum is only about 7% of the total brain mass, while the killer whale cerebellum is almost 14%. These results are significant because they contradict claims that the cerebellum scales proportionally with the rest of the brain in all mammals. They also correct the generalization that all cetaceans have enlarged cerebella. We suggest possible reasons for the existence of such a large cerebellar size difference between these two species. Cerebellar function is not fully understood, and comparing the abilities of animals with differently sized cerebella can help uncover functional roles of the cerebellum in humans and animals. Here we show that the large cerebellar difference likely relates to evolutionary history, diving, sensory capability, and ecology.

Apneustic anesthesia ventilation improves pulmonary function in anesthetized bottlenose dolphins (Tursiops truncatus).

Introduction: Use of mechanical ventilation during general anesthesia is a necessary practice in the anesthetization of small cetaceans as spontaneous ventilation fails to provide adequate gas exchange. Currently available methods of ventilation do not account for the intermittent breathing strategy of representative species within this infraorder of fully aquatic mammals and may have a significant effect on cardiac and respiratory physiology. Methods: To understand the impact of mechanical ventilation on cardiopulmonary function in one small species of cetacean, the bottlenose dolphin (Tursiops truncatus), we compared controlled mechanical ventilation (CMV) to a novel ventilation method known as apneustic anesthesia ventilation (AAV). AAV simulates the normal inspiratory breath-hold pattern of dolphins. Ten anesthetic procedures (dental procedure, n = 9; bronchoscopy, n = 2) were performed on nine dolphins (age range: 10-42 years; mean = 32 years; median = 37 years; female = 3, 40%; male = 6, 60%). In a cross-over study design, dolphins were instrumented and randomly assigned to AAV or CMV as the initial mode of ventilation, then switched to the alternate mode. Baseline cardiopulmonary data were collected and again after 30 min on each mode of ventilation. Cardiac index, stroke volume index, systemic vascular resistance, alveolar dead space, alveolar-arterial oxygen tension gradient, arterial oxygen content, oxygen delivery index, and dynamic respiratory system compliance index were calculated at each of the four time points. Results: During AAV, dolphins had higher arterial oxygen tension, higher mean airway pressure, reduced alveolar dead space ventilation and lower alveolar-arterial oxygen difference. Cardiovascular performance was not statistically different between the two modes. Discussion: Our study suggests AAV, which more closely resembles the conscious intermittent respiratory pattern phenotype of dolphins, improves ventilation and pulmonary function in the anesthetized dolphin. Future studies should evaluate the cardiopulmonary effects of neutral buoyancy and cardiopulmonary sparing drug protocols to reduce the need for hemodynamic support of current protocols.

Diagnosis and successful treatment of a lung abscess associated with Brucella species infection in a bottlenose dolphin (Tursiops truncatus).

This brief communication describes the clinical presentation, antemortem diagnosis, and successful treatment of a pulmonary abscess associated with a Brucella sp. in a 27-yr-old female bottlenose dolphin (Tursiops truncatus). Ultrasound revealed a 3-cm diameter hypoechoic mass deep to the pleural lining in the left lung field. Multiple ultrasound-guided fine-needle aspirates were performed and tested for bacterial and fungal etiology. All cultures were negative, but the infectious agent was identified by MicroSEQ analysis in two samples and confirmed with real-time polymerase chain reaction (PCR) amplification using known Brucella sp. primers. Amikacin was infused into the abscess and was followed by an oral doxycycline and rifampin protocol. Follow-up diagnostic imaging, including radiographs and computed tomography, revealed a resolved lesion with minimal mineralization within the affected lung fields. Brucellosis should be considered for pulmonary disease in dolphins, and personnel who interact with marine animals should use caution to prevent zoonotic brucellosis.

Sights and sounds dolphins, Tursiops truncatus preying on native fish of San Diego Bay and offshore in the Pacific Ocean.

For the first time, dolphins wearing video cameras were observed capturing and eating live native fish. While freely swimming in San Diego Bay, one dolphin caught 69 resident fish, 64 demersal, 5 near surface, while the other caught 40, 36 demersal and 4 near the surface. Two other dolphins were observed capturing 135 live native fish in a sea water pool. Two additional dolphins were observed feeding opportunistically during open water sessions in the Pacific Ocean. Notably, one of these dolphins was observed to consume 8 yellow-bellied sea snakes (Hydrophis platurus). Searching dolphins clicked at intervals of 20 to 50 ms. On approaching prey, click intervals shorten into a terminal buzz and then a squeal. Squeals were bursts of clicks that varied in duration, peak frequency, and amplitude. Squeals continued as the dolphin seized, manipulated and swallowed the prey. If fish escaped, the dolphin continued the chase and sonar clicks were heard less often than the continuous terminal buzz and squeal. During captures, the dolphins' lips flared to reveal nearly all of the teeth. The throat expanded outward. Fish continued escape swimming even as they entered the dolphins' mouth, yet the dolphin appeared to suck the fish right down.

Signature whistles exhibit a 'fade-in' and then 'fade-out' pattern of relative amplitude declination.

Bottlenose dolphins have individually distinct signature whistles that are characterized by a stereotyped frequency-time contour. Signature whistles are commonly exchanged with short time delays between calls. Dolphin whistles are produced by pressurized nasal sacs that increase and then decrease in pressure over emission. This study found that the relative amplitude modulation pattern over time exhibited the same fade-in and then fade-out pattern in the signature whistles of eight bottlenose dolphins at the Navy in San Diego, CA. Both the initial and final five percent of the whistle's duration also had significantly lower mean relative amplitude than the center five percent. The current analyses of the amplitude-time relationship was then integrated to a previously reported model of the negative relationship between relative log amplitude and log peak frequency. This produced a more robust model for accounting for the predictable aspects of the more broadly non-stereotyped amplitude modulations of signature whistles. Whether dolphins can intentionally manipulate these amplitude features or they are simple by-products of the sound production system, and further whether they are perceived and utilized by receivers, is an exciting area for continued research.

Non-stereotyped amplitude modulation across signature whistle contours.

Bottlenose dolphin signature whistles are characterized by distinctive frequency modulation over time. The stable frequency contours of these whistles broadcast individual identity information. Little is known however, about whether or not the amplitude contour is also stereotyped. Here, we examined the relative amplitude-time contour of signature whistle emissions from eight bottlenose dolphins (Tursiops truncatus) in the U.S. Navy Marine Mammal Program (MMP) in San Diego, CA. The results suggested that unlike the stable frequency-time contour, the amplitude-time contour of signature whistles were largely non-stereotyped, characterized by large variability across multiple whistle emissions. Relative amplitude was negatively related to log peak frequency, with more energy focused in the lower frequency bands. This trend was consistent over all eight dolphins despite having quite different signature whistle contours. This relationship led to the amplitude contours being slightly more stereotyped within than between dolphins. We propose that amplitude across signature whistle emissions may serve as an avenue for encoding additional communicative information. We encourage future studies to incorporate analyses of amplitude contours in addition to frequency contours of signature whistles in order to begin to understand what role it may play in the dolphin communication system.

Dolphins as animal models for type 2 diabetes: sustained, post-prandial hyperglycemia and hyperinsulinemia.

There is currently no known natural animal model that fully complements type 2 diabetes in humans. Criteria for a true natural animal model include the presence of a fasting hyperglycemia, evidence of insulin resistance, and pathologies matching that reported in humans. To investigate the bottlenose dolphin (Tursiops truncatus) as a comparative model for type 2 diabetes in humans, hourly plasma and urine chemistry changes, including glucose, were analyzed among five healthy, adult dolphins for 24 h following ingestion of 2.5-3.5 kg of mackerel or 2-3 L of 10% dextrose in ionosol. Fasting and 2 h post-prandial insulin levels were also determined among five adult dolphins to assess the presence of hyperinsulinemia. Finally, a case-control study compared insulin and glucagon levels among dolphins with and without iron overload, a condition associated with insulin resistance in humans. Both protein and dextrose meals caused significant increases in plasma glucose during the 0-5 h post-prandial period; dolphins fed dextrose demonstrated a sustained hyperglycemia lasting 5-10 h. Fasting plasma insulin levels among healthy dolphins mimicked those found in humans with some insulin resistance. Dolphins with hemochromatosis had higher post-prandial plasma insulin levels compared to controls. We conclude that bottlenose dolphins can demonstrate metabolic responses consistent with type 2 diabetes, specifically sustained hyperglycemia and hyperinsulinemia. Understanding more about how and why dolphins have a diabetes-like metabolism may provide new research avenues for diabetes in humans.

Neural time and movement time in choice of whistle or pulse burst responses to different auditory stimuli by dolphins.

Echolocating dolphins emit trains of clicks and receive echoes from ocean targets. They often emit each successive ranging click about 20 ms after arrival of the target echo. In echolocation, decisions must be made about the target--fish or fowl, predator or food. In the first test of dolphin auditory decision speed, three bottlenose dolphins (Tursiops truncatus) chose whistle or pulse burst responses to different auditory stimuli randomly presented without warning in rapid succession under computer control. The animals were trained to hold pressure catheters in the nasal cavity so that pressure increases required for sound production could be used to split response time (RT) into neural time and movement time. Mean RT in the youngest and fastest dolphin ranged from 175 to 213 ms when responding to tones and from 213 to 275 ms responding to pulse trains. The fastest neural times and movement times were around 60 ms. The results suggest that echolocating dolphins tune to a rhythm so that succeeding pulses in a train are produced about 20 ms over target round-trip travel time. The dolphin nervous system has evolved for rapid processing of acoustic stimuli to accommodate for the more rapid sound speed in water compared to air.

Evaluation of population health among bottlenose dolphins (Tursiops truncatus) at the United States Navy Marine Mammal Program.

OBJECTIVE: To evaluate health indicators for a population of bottlenose dolphins in the US Navy Marine Mammal Program (MMP) by use of data acquired from 1988 through 2007. DESIGN: Retrospective cohort study. ANIMALS: 167 bottlenose dolphins. PROCEDURES: The following indicators were used to evaluate the health of dolphins during the 20-year period: 5-year age structure, median survival age, annual survival rates, mortality rates, and neonatal and calf survival and mortality rates. Limitations of these population measurements as health indicators for dolphins were assessed. RESULTS: Crude mortality rates of dolphins for 1988 through 1992, 1993 through 1997, 1998 through 2002, and 2003 through 2007 were 3.1%, 4.7%, 3.6%, and 2.4%, respectively; during these same 4 study periods, median survival ages were 14.3, 14.4, 17.7, and 26.1 years, respectively, and mean survival rates were 0.98, 0.97, 0.97, and 0.99, respectively. From 1988 through 1997, 1998 through 2002, and 2003 through 2007, neonatal mortality rates were 4 of 16, 5 of 20, and 2 of 14 neonates, respectively. During these 3 study periods, mean annual survival rates for calves < 3 years old (excluding neonates that died at < 30 days old) were 0.97, 0.92, and 0.99, respectively. CONCLUSIONS AND CLINICAL RELEVANCE: Although there were limitations to the measurement of some health indicators, use of multiple methods indicated that the health of dolphins in the MMP population was comparable to, if not better than, that published for other dolphin populations. The MMP population of dolphins may provide useful reference values of health indicators for use in assessment of other managed dolphin populations.

A method to enable a bottlenose dolphin (Tursiops truncatus) to echolocate while out of water.

The study of site-specific brain activity associated with dolphin echolocation has been hampered by the difficulties inherent in administering radiolabels and performing medical imaging while a dolphin echolocates in an aquatic environment. To overcome these limitations, a system has been developed to allow a bottlenose dolphin to echolocate while out of the water. The system relies on a "phantom echo generator" (PEG) consisting of a Texas Instruments C6713 digital signal processor with an analog input/output daughtercard. Echolocation clicks produced by the dolphin are detected with a hydrophone embedded in a suction cup on the melon, then digitized within the PEG. Clicks exceeding a user-defined threshold are convolved with a target impulse response, delayed, and scaled before being converted to analog and transmitted through a sound projector embedded in a suction cup attached to the dolphin's lower jaw. Dolphin in-air echolocation behavior, inter-click intervals, and overall performance were analogous to those observed during comparable underwater testing with physical targets, demonstrating that the dolphin was indeed performing an echolocation task while out of water.

Relationship of blood flow and metabolism to acoustic processing centers of the dolphin brain.

Odontocete brain tissues associated with auditory processing are hypertrophied and modified relative to their terrestrial counterparts. The relationship between the functional demand on these tissues and metabolic substrate requirements is unknown. Using positron emission tomography (PET), relative cerebral blood flow was measured in a bottlenose dolphin. Approximately 60 mCi (13)NH(3) was administered to the dolphin via a catheter inserted into the hepatic vein and threaded proximate to the vena cava. Radiolabel initially appeared as distributed focal points in the cerebellum. Increasing scan time resulted in an increase in the number of focal regions and in the diffusivity of label activity throughout the brain. The time course and spatial distribution of radiolabel was consistent with a cerebral blood supply dominated by the spinal meningeal arteries. Blood flow was predominantly observed in the cerebellum and neocortex, particularly the auditory and visual cortex. Differential brain glucose uptake, previously measured in a separate dolphin, showed good agreement with the differential supply of blood to brain tissues. Rates of blood supply and glucose uptake in the auditory cortex, inferior colliculus, and cerebellum are consistent with a high metabolic demand of tissues which are important to the integration of auditory and other sensory inputs.

Five members of a mixed-sex group of bottlenose dolphins share a stereotyped whistle contour in addition to maintaining their individually distinctive signature whistles.

Most commonly, animal communication systems are driven by shared call repertoires, with some individual distinctiveness encoded as a byproduct of voice cues. We provide evidence that bottlenose dolphins produce both individually distinctive whistles, and a shared whistle type. A stereotyped whistle contour (termed the group whistle) is shared by five bottlenose dolphins that have lived, worked, and traveled together for at least 21 years. These five dolphins are members of a group of eight dolphins that work as a specialized team for the Navy Marine Mammal Program. Each dolphin is routinely recorded during periods when an individual is isolated from the others in above ground pools as part of their routine training. Each of the eight dolphins has an individually distinctive signature whistle. In addition, at least five of these dolphins share a distinct non-signature whistle type. This shared whistle contour was produced an average of 22.4% +/- 9.0% of the time during periods in which individuals were isolated. During these isolations the signature whistle was produced an average of 42.9% +/- 11.9% of the time. This is consistent with decades of signature whistle research. A group of 10 naïve observers rated the similarity of the different whistle contours. The observers rated the group whistle contour produced by all five dolphins as highly similar (P < 0.01). Their ratings further showed that the signature whistles of the five dolphins were very different (P < 0.01). These findings were further supported by discriminant function analyses. That said, the shared whistle contours still exhibited individual differences which may allow conspecifics to identify the producer even when a whistle contour is shared among multiple dolphins. This is the first in-depth analysis of a non-signature whistle type shared among multiple conspecifics.

Use of phlebotomy treatment in Atlantic bottlenose dolphins with iron overload.

CASE DESCRIPTION: 3 adult (24- to 43-year-old) Atlantic bottlenose dolphins (Tursiops truncatus) with chronic episodic malaise and inappetence associated with high serum aminotransferase (alanine aminotransferase and aspartate aminotransferase) activities, high serum iron concentration, and serum transferrin saturation > 80% were evaluated. CLINICAL FINDINGS: Results of histologic examination of liver biopsy specimens revealed hemosiderosis in all 3 dolphins. Except for chronic lymphocytosis in 1 dolphin, results of extensive diagnostic testing revealed no other abnormalities. For each dolphin, a diagnosis of iron overload of unknown origin was made. TREATMENT AND OUTCOME: Phlebotomy treatment was implemented to reduce body stores of iron. Each phlebotomy procedure removed 7% to 17% (1 to 3 L) of estimated blood volume. Treatment consisted of an induction phase of weekly phlebotomy procedures for 22 to 30 weeks, which was complete when serum iron concentration and aminotransferase activities were within reference ranges and serum transferrin saturation was < or = 20% or Hct was < or = 30%. Total amount of iron removed from each dolphin was 53 to 111 mg/kg (24.1 to 50.5 mg/lb) of body weight. One dolphin required maintenance procedures at 8- to 12-week intervals when high serum iron concentration was detected. CLINICAL RELEVANCE: Although the cause of the iron overload and high serum aminotransferase activities remained unknown, phlebotomy treatment successfully resolved the clinicopathologic abnormalities, supporting a role of iron overload in the hepatopathy of the 3 dolphins.

Higher neuron densities in the cerebral cortex and larger cerebellums may limit dive times of delphinids compared to deep-diving toothed whales.

Since the work of Tower in the 1950s, we have come to expect lower neuron density in the cerebral cortex of larger brains. We studied dolphin brains varying from 783 to 6215g. As expected, average neuron density in four areas of cortex decreased from the smallest to the largest brain. Despite having a lower neuron density than smaller dolphins, the killer whale has more gray matter and more cortical neurons than any mammal, including humans. To begin a study of non-dolphin toothed whales, we measured a 596g brain of a pygmy sperm whale and a 2004g brain of a Cuvier's beaked whale. We compared neuron density of Nissl stained cortex of these two brains with those of the dolphins. Non-dolphin brains had lower neuron densities compared to all of the dolphins, even the 6215g brain. The beaked whale and pygmy sperm whale we studied dive deeper and for much longer periods than the dolphins. For example, the beaked whale may dive for more than an hour, and the pygmy sperm whale more than a half hour. In contrast, the dolphins we studied limit dives to five or 10 minutes. Brain metabolism may be one feature limiting dolphin dives. The brain consumes an oversized share of oxygen available to the body. The most oxygen is used by the cortex and cerebellar gray matter. The dolphins have larger brains, larger cerebellums, and greater numbers of cortex neurons than would be expected given their body size. Smaller brains, smaller cerebellums and fewer cortical neurons potentially allow the beaked whale and pygmy sperm whale to dive longer and deeper than the dolphins. Although more gray matter, more neurons, and a larger cerebellum may limit dolphins to shorter, shallower dives, these features must give them some advantage. For example, they may be able to catch more elusive individual high-calorie prey in the upper ocean.

Cetacean sleep: an unusual form of mammalian sleep.

Our knowledge of the form of lateralized sleep behavior, known as unihemispheric slow wave sleep (USWS), seen in all members of the order Cetacea examined to date, is described. We trace the discovery of this phenotypically unusual form of mammalian sleep and highlight specific aspects that are different from sleep in terrestrial mammals. We find that for cetaceans sleep is characterized by USWS, a negligible amount or complete absence of rapid eye movement (REM) sleep, and a varying degree of movement during sleep associated with body size, and an asymmetrical eye state. We then compare the anatomy of the mammalian somnogenic system with what is known in cetaceans, highlighting areas where additional knowledge is needed to understand cetacean sleep. Three suggested functions of USWS (facilitation of movement, more efficient sensory processing and control of breathing) are discussed. Lastly, the possible selection pressures leading to this form of sleep are examined, leading us to the suggestion that the selection pressure necessitating the evolution of cetacean sleep was most likely the need to offset heat loss to the water from birth and throughout life. Aspects such as sentinel functions and breathing are likely to be proximate evolutionary phenomenon of this form of sleep.