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Odontocetes have evolved for millions of years to own a unique echolocation system. The exceptional performance of odontocetes echolocation system can provide reference to artificial sonar systems, acoustic metamaterials and sound control designs. Research on odontocetes biosonar requires interdisciplinary effort, including acoustics, biology, biomimetics, anatomy, physiology and signal analysis. In this paper, we review odontoctes’ biosonar emission process from aspects of anatomy, biosonar signal and beam formation. To begin, computed tomography scanning and untrasound measurements are combined to reconstruct the sound speed and density distributions. To follow, efforts are thrown to probe into the biosonar signal and its corresponding acoustic behavior. Numerical simulations are used to investigate the odontocetes’ biosonar beam formation. The secret of exceptional performance of odontocetes’ echolocation system lies in their unique anatomy. Odontocete integrates acoustic structures with different acoustic impedances, namely solid bony structures, air space and soft tissues as a whole emission system to efficiently modulate sound propagation and sound beam formation. These acoustic structures are well organized in the forehead, forming a natural acoustic metamaterial to perform a good control of sounds. These results can enlighten artificial sonar designs.
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Keywords:
- odontocetes /
- echolocation /
- biosonar /
- computed tomography scan
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图 1 (a) 东亚江豚头部的三维重建; (b) 小抹香鲸头部的三维重建; (c) 中华白海豚头部的三维重建
Figure 1. (a) Three-dimensional reconstruction of a finless porpoise head; (b) three-dimensional reconstruction of a pygmy sperm whale head; (c) three-dimensional reconstruction of an Indo-Pacific humpback dolphin.
图 2 (a) 东亚江豚头部声学结构的三维重建; (b)小抹香鲸头部声学结构的三维重建; (c) 中华白海豚头部声学结构的三维重建结果; 青色、棕色、红色、黄色和灰色分别表示结缔组织、额隆、声源、气囊与上颌骨
Figure 2. (a) Three-dimensional acoustic structure reconstructions of a finless porpoise; (b) three-dimensional acoustic structure reconstructions of a pygmy sperm whale; (c) three-dimensional acoustic structure reconstructions of an Indo-Pacific humpback dolphin. Upper jaw, sound source, air components (including the air sacs and nasal passage) are represented in different colors.
图 3 (a)东亚江豚头部声发射系统的水平截面声速重建; (b)东亚江豚头部声发射系统的垂直截面声速重建[34]; (c)小抹香鲸头部声发射系统的水平截面声速重建; (d)小抹香鲸头部声发射系统的垂直截面声速重建[36]; (e)中华白海豚头部声发射系统的水平截面声速重建; (f)中华白海豚头部声发射系统的垂直截面声速重建[38]
Figure 3. (a) Sound speed reconstructions of finless porpoise for sound emission system in horizontal section; (b) sound speed reconstructions of finless porpoise for sound emission system in vertical section[34]; (c) sound speed reconstructions of pygmy sperm whale for sound emission system in horizontal section; (d) sound speed reconstructions of pygmy sperm whale for sound emission system in vertical section[36]; (e) sound speed reconstructions of the Indo-Pacific humpback dolphin for sound emission system in horizontal section; (f) sound speed reconstructions of the Indo-Pacific humpback dolphin for sound emission system in vertical section[38].
图 5 (a)中华白海豚不同能量回声定位脉冲的频谱分布; (b) 东亚江豚不同能量回声定位脉冲信号的频谱分布; 图中的–3 dB, –6 dB, –10 dB以及–20 dB分别表示能量处于最高能量声信号–3 dB到0 dB范围, –6 dB到–3 dB范围, –10 dB到–6 dB范围以及–20 dB到–10 dB范围的回声定位信号
Figure 5. (a) Mean spectrum of the clicks from –3 dB, –6 dB, and –10 dB groups for the Indo-Pacific humpback dolphin; (b) the mean spectrum of the clicks from –3 dB, –6 dB, –10 dB and –20 dB groups for the finless porpoise.
图 6 (a)宽吻海豚固定频率型通信声信号时频图; (b)宽吻海豚正弦型通信声信号时频图; (c)宽吻海豚凸型通信声信号时频图; (d)宽吻海豚凹型通信声信号时频图; (e) 宽吻海豚上扫频型通信声信号时频图; (f) 宽吻海豚下扫频型通信声信号时频图[58,59]; 其中颜色深浅表示声信号强度大小
Figure 6. (a) Spectrogram of constant frequency whistles of the bottlenose dolphins; (b) the spectrogram sinusoidal whistles of the bottlenose dolphins; (c) the spectrogram of convex or hill whistles of the bottlenose dolphins; (d) the spectrogram of concave or valley whistles of the bottlenose dolphins; (e) the spectrogram of upsweep whistles of the bottlenose dolphins; (f) the spectrogram of down sweep frequency whistles of the bottlenose dolphins[58,59].
图 7 (a) 厦门五缘湾圈养两只宽吻海豚处在自由游动状态; (b) 宽吻海豚处于训练状态; (c)宽吻海豚处于自由游动时发出的通信声信号类别及其占比; (d) 宽吻海豚处于训练状态下发出的通信信号的类别及其占比[58,59]
Figure 7. (a) Two captive free swimming bottlenose dolphins in Xiamen; (b) bottlenose dolphins under training; (c) pie chart of the classified whistles of two bottlenose dolphins under free swimming; (d) pie chart of the classified whistles of two bottlenose dolphins under training conditions[58,59].
图 8 (a) 中心频率为130 kHz的声脉冲经过鼠海豚无头骨模型调控形成的声波波束; (b) 中心频率为130 kHz的声脉冲经过鼠海豚完整模型调控形成的声波波束[31]
Figure 8. (a) Beam directivity of a sound pulse with a centroid frequency of 130 kHz for No-Skull model of harbor porpoise; (b) the beam directivity of a sound pulse with a centroid frequency of 130 kHz for a complete model of harbor porpoise[31].
图 9 (a) 声波在白鱀豚垂直截面第一传播时刻的声场分布; (b) 声波在白鱀豚垂直截面第二传播时刻的声场分布, 其中RW与IW分别表示传播过程中的反射波与表面波; (c) 声波在白鱀豚垂直截面第三传播时刻的声场分布; (d)第一传播时刻声场分布放大; (e)第二传播时刻声场分布放大; (f)第三传播时刻声场分布放大[74]
Figure 9. (a) Propagation plot of a short-duration impulse source for Baiji in vertical section at time 1; (b) propagation plot of a short-duration impulse source for Baiji in vertical section at time 2; (c) propagation plot of a short-duration impulse source for Baiji in vertical section at time 3; (d) enlarged details of (a); (e) enlarged details of (b); (f) enlarged details of (c)[74].
图 11 (a) 声波在小抹香鲸完整头部模型第一传播时刻的声场分布; (b) 声波在小抹香鲸完整头部模型第二传播时刻的声场分布; (c) 声波在小抹香鲸完整头部模型第三传播时刻的声场分布; (d)声波在小抹香鲸头部无气体结构模型第一传播时刻的声场分布; (e) 声波在小抹香鲸头部无气体结构模型第二传播时刻的声场分布; (f) 声波在小抹香鲸头部无气体结构模型第三传播时刻的声场分布; (g) 峰值频率为125 kHz的声脉冲经过小抹香鲸头部完整模型调控形成的声波波束; (h) 峰值频率为125 kHz的声脉冲经过小抹香鲸头部无气体结构模型调控形成的声波波束[35]
Figure 11. (a) Propagation plot of the transient sound waves at time 1 under a full model case of pygmy sperm whale; (b) propagation plot of the transient sound waves at time 2 under a full model case of pygmy sperm whale; (c) propagation plot of the transient sound waves at time 3 under a full model case of pygmy sperm whale; (d) propagation plot of the transient sound waves at time 1 under a model case without air components; (e) propagation plot of the transient sound waves at time 2 under a model case without air components; (f) propagation plot of the transient sound waves at time 3 under a model case without air components; (g) the beam directivity of a sound pulse with a peak frequency of 125 kHz for the full model case of the pygmy sperm whale; (h) the beam directivity of a sound pulse with a peak frequency of 125 kHz for the model case of pygmy sperm whale without air components[35].
图 12 (a)鼠海豚无额隆模型和完整模型在第一传播时刻的传播声场分布; (b)鼠海豚无额隆模型和完整模型在第二传播时刻的声场分布; (c) 鼠海豚无额隆模型和完整模型在第三传播时刻的声场分布; (d) 鼠海豚无额隆模型和完整模型在第四传播时刻的声场分布; (e) 中心频率为130 kHz的声脉冲经过鼠海豚无额隆模型调控形成的声波波束; (f) 中心频率为130 kHz的声脉冲经过鼠海豚完整模型调控形成的声波波束[31]
Figure 12. (a) Acoustic field of no-melon and full head cases at time 1; (b) acoustic field of no-melon and full head cases at time 2; (c) acoustic field of no-melon and full head cases at time 3; (d) acoustic field of no-melon and full head cases at time 4; (e) the beam directivity of a sound pulse with a centroid frequency of 130 kHz for no-melon case of harbor porpoise; (f) the beam directivity of a sound pulse with a centroid frequency of 130 kHz for the full head case[31].
图 13 (a) 头部压缩对峰值频率为125 kHz的声脉冲传播形成的声场的影响, 其中θ表示的是前庭囊的倾斜角, NA代表的是相对于原始模型的归一化面积; (b) 五种模型相应的波束分布特性; (c) 五种模型形成的声波波束的–3 dB带宽与主瓣角分布[34]
Figure 13. (a) Compressing effect of models I, II, III, IV, and V on acoustic field of the sound pulse with a peak frequency of 125 kHz inside the head, where θ represents the orientation angle of the vestibular sac and NA represents the normalized area of the forehead tissues with respect to those of the original model I; (b) beam directivities of the five cases; (c) sound beams’ –3 dB beam widths and main beam angle distribution of the five cases[34].
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