1Sound is the primary sensory cue for most marine mammals, and this is especially true for 2 cetaceans. To passively and actively acquire information about their environment, cetaceans 3 have perhaps the most derived ears of all mammals, capable of sophisticated, sensitive hearing 4 and auditory processing. These capabilities have developed for survival in an underwater world 5 where sound travels five times faster than in air, and where light is quickly attenuated and often 6 limited at depth, at night, and in murky waters. Cetacean auditory evolution has capitalized on 7 the ubiquity of sound cues and the efficiency of underwater acoustic communication. The sense 8 of hearing is central to cetacean sensory ecology, enabling vital behaviors such as locating prey, 9 detecting predators, identifying conspecifics, and navigating. Increasing levels of anthropogenic 10 ocean noise appears to influence many of these activities. 11Here we describe the historical progress of investigations on cetacean hearing, with a 12 particular focus on odontocetes and recent advancements. While this broad topic has been 13 studied for several centuries, new technologies in the last two decades have been leveraged to 14 improve our understanding of a wide range of taxa, including some of the most elusive species. 15 This paper addresses topics including how sounds are received, what sounds are detected, 16 hearing mechanisms for complex acoustic scenes, recent anatomy and physiology studies, the 17 potential impacts of noise, and mysticete hearing. We conclude by identifying emerging 18 research topics and areas which require greater focus. 19 20 3
Cetaceans possess highly derived auditory systems adapted for underwater hearing. Odontoceti (toothed whales) are thought to receive sound through specialized fat bodies that contact the tympanoperiotic complex, the bones housing the middle and inner ears. However, sound reception pathways remain unknown in Mysticeti (baleen whales), which have very different cranial anatomies compared to odontocetes. Here, we report a potential fatty sound reception pathway in the minke whale (Balaenoptera acutorostrata), a mysticete of the balaenopterid family. The cephalic anatomy of seven minke whales was investigated using computerized tomography and magnetic resonance imaging, verified through dissections. Findings include a large, well-formed fat body lateral, dorsal, and posterior to the mandibular ramus and lateral to the tympanoperiotic complex. This fat body inserts into the tympanoperiotic complex at the lateral aperture between the tympanic and periotic bones and is in contact with the ossicles. There is also a second, smaller body of fat found within the tympanic bone, which contacts the ossicles as well. This is the first analysis of these fatty tissues' association with the auditory structures in a mysticete, providing anatomical evidence that fatty sound reception pathways may not be a unique feature of odontocete cetaceans. Anat Rec, 2012. © 2012 Wiley Periodicals, Inc.
Whales receive underwater sounds through a fundamentally different mechanism than their close terrestrial relatives. Instead of hearing through the ear canal, cetaceans hear through specialized fatty tissues leading to an evolutionarily novel feature: an acoustic funnel located anterior to the tympanic aperture. We traced the ontogenetic development of this feature in 56 fetal specimens from 10 different families of toothed (odontocete) and baleen (mysticete) whales, using X-ray computed tomography. We also charted ear ossification patterns through ontogeny to understand the impact of heterochronic developmental processes. We determined that the acoustic funnel arises from a prominent V-shaped structure established early in ontogeny, formed by the malleus and the goniale. In odontocetes, this V-formation develops into a cone-shaped funnel facing anteriorly, directly into intramandibular acoustic fats, which is likely functionally linked to the anterior orientation of sound reception in echolocation. In contrast, the acoustic funnel in balaenopterids rotates laterally, later in fetal development, consistent with a lateral sound reception pathway. Balaenids and several fossil mysticetes retain a somewhat anteriorly oriented acoustic funnel in the mature condition, indicating that a lateral sound reception pathway in balaenopterids may be a recent evolutionary innovation linked to specialized feeding modes, such as lunge-feeding.
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Like elephants, baleen whales produce low-frequency (LF) and even infrasonic (IF) signals, suggesting they may be particularly susceptible to underwater anthropogenic sound impacts. Analyses of computerized tomography scans and histologies of the ears in five baleen whale and two elephant species revealed that LF thresholds correlate with basilar membrane thickness/width and cochlear radii ratios. These factors are consistent with high-mass, low-stiffness membranes and broad spiral curvatures, suggesting that Mysticeti and Proboscidea evolved common inner ear adaptations over similar time scales for processing IF/LF sounds despite operating in different media.
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