Mechanical properties of a bio-inspired robotic knifefish with an undulatory propulsor

South American electric knifefish are a leading model system within neurobiology. Recent efforts have focused on understanding their biomechanics and relating this to their neural processing strategies. Knifefish swim by means of an undulatory fin that runs most of the length of their body, affixed to the belly. Propelling themselves with this fin enables them to keep their body relatively straight while swimming, enabling straightforward robotic implementation with a rigid hull. In this study, we examined the basic properties of undulatory swimming through use of a robot that was similar in some key respects to the knifefish. As we varied critical fin kinematic variables such as frequency, amplitude, and wavelength of sinusoidal traveling waves, we measured the force generated by the robot when it swam against a stationary sensor, and its velocity while swimming freely within a flow tunnel system. Our results show that there is an optimal operational region in the fin's kinematic parameter space. The optimal actuation parameters found for the robotic knifefish are similar to previously observed parameters for the black ghost knifefish, Apteronotus albifrons. Finally, we used our experimental results to show how the force generated by the robotic fin can be decomposed into thrust and drag terms. Our findings are useful for future bio-inspired underwater vehicles as well as for understanding the mechanics of knifefish swimming.     

Center of mass motion in swimming fish: effects of speed and locomotor mode during undulatory propulsion

Studies of center of mass (COM) motion are fundamental to understanding the dynamics of animal movement, and have been carried out extensively for terrestrial and aerial locomotion. But despite a large amount of literature describing different body movement patterns in fishes, analyses of how the center of mass moves during undulatory propulsion are not available. These data would be valuable for understanding the dynamics of different body movement patterns and the effect of differing body shapes on locomotor force production. In the present study, we analyzed the magnitude and frequency components of COM motion in three dimensions (x: surge, y: sway, z: heave) in three fish species (eel, bluegill sunfish, and clown knifefish) swimming with four locomotor modes at three speeds using high-speed video, and used an image cross-correlation technique to estimate COM motion, thus enabling untethered and unrestrained locomotion. Anguilliform swimming by eels shows reduced COM surge oscillation magnitude relative to carangiform swimming, but not compared to knifefish using a gymnotiform locomotor style. Labriform swimming (bluegill at 0.5 body lengths/s) displays reduced COM sway oscillation relative to swimming in a carangiform style at higher speeds. Oscillation frequency of the COM in the surge direction occurs at twice the tail beat frequency for carangiform and anguilliform swimming, but at the same frequency as the tail beat for gymnotiform locomotion in clown knifefish. Scaling analysis of COM heave oscillation for terrestrial locomotion suggests that COM heave motion scales with positive allometry, and that fish have relatively low COM oscillations for their body size.     

Musculoskeletal morphology and regionalization within the dorsal and anal fins of bluegill sunfish (Lepomis macrochirus)

Ray‐finned fishes actively control the shape and orientation of their fins to either generate or resist hydrodynamic forces. Because of the emergent mechanical properties of their segmented, bilaminar fin rays (lepidotrichia), and actuation by multiple muscles, fish can control the rigidity and curvature of individual rays independently, thereby varying the resultant forces across the fin surfaces. Expecting that differences in fin‐ray morphology should reflect variation in their mechanical properties, we measured several musculoskeletal features of individual spines and rays of the dorsal and anal fins of bluegill sunfish, Lepomis macrochirus, and assessed their mobility and flexibility. We separated the fin‐rays into four groups based on the fin (dorsal or anal) or fin‐ray type (spine or ray) and measured the length of the spines/rays and the mass of the three median fin‐ray muscles: the inclinators, erectors and depressors. Within the two ray groups, we measured the portion of the rays that were segmented vs. unsegmented and branched vs. unbranched. For the majority of variables tested, we found that variations between fin‐rays within each group were significantly related to position within the fin and these patterns were conserved between the dorsal and anal rays. Based on positional variations in fin‐ray and muscle parameters, we suggest that anterior and posterior regions of each fin perform different functions when interacting with the surrounding fluid. Specifically, we suggest that the stiffer anterior rays of the soft dorsal and anal fins maintain stability and keep the flow across the fins steady. The posterior rays, which are more flexible with a greater range of motion, fine‐tune their stiffness and orientation, directing the resultant flow to generate lateral and some thrust forces, thus acting as an accessory caudal fin. J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

Functional morphology of the fin rays of teleost fishes

Ray‐finned fishes are notable for having flexible fins that allow for the control of fluid forces. A number of studies have addressed the muscular control, kinematics, and hydrodynamics of flexible fins, but little work has investigated just how flexible ray‐finned fish fin rays are, and how flexibility affects their response to environmental perturbations. Analysis of pectoral fin rays of bluegill sunfish showed that the more proximal portion of the fin ray is unsegmented while the distal 60% of the fin ray is segmented. We examined the range of motion and curvatures of the pectoral fin rays of bluegill sunfish during steady swimming, turning maneuvers, and hovering behaviors and during a vortex perturbation impacting the fin during the fin beat. Under normal swimming conditions, curvatures did not exceed 0.029 mm^?1 in the proximal, unsegmented portion of the fin ray and 0.065 mm^?1 in the distal, segmented portion of the fin ray. When perturbed by a vortex jet traveling at approximately 1 ms^?1 (67 ± 2.3 mN s.e. of force at impact), the fin ray underwent a maximum curvature of 9.38 mm^?1. Buckling of the fin ray was constrained to the area of impact and did not disrupt the motion of the pectoral fin during swimming. Flexural stiffness of the fin ray was calculated to be 565 × 10^?6 Nm². In computational fluid dynamic simulations of the fin‐vortex interaction, very flexible fin rays showed a combination of attraction and repulsion to impacting vortex dipoles. Due to their small bending rigidity (or flexural stiffness), impacting vortices transferred little force to the fin ray. Conversely, stiffer fin rays experienced rapid small‐amplitude oscillations from vortex impacts, with large impact forces all along the length of the fin ray. Segmentation is a key design feature of ray‐finned fish fin rays, and may serve as a means of making a flexible fin ray out of a rigid material (bone). This flexibility may offer intrinsic damping of environmental fluid perturbations encountered by swimming fish. J. Morphol. 274:1044–1059, 2013. © 2013 Wiley Periodicals, Inc.     

Better Endurance and Load Capacity: An Improved Design of Manta Ray Robot (RoMan-II)

An improved design of a biomimetic underwater vehicle (RoMan-II) inspired by manta ray is presented in this paper. The design of the prototype and the swimming motion control are discussed. Instead of using rigid multiple degree-of-freedom linkages as fin rays in the first version, six flexible fin rays are adopted to drive two sided fins which generate thrust through flapping motions. Furthermore, in order to save the energy for a long distance cruising, a bio-inspired gliding motion is incorporated onto the motion control of the improved prototype. With a closed-loop buoyancy control system, the vehicle can perform gliding locomotion in water, which reduces the overall energy consumption. The vehicle can also perform pivot turning and backward locomotion without turning its body. It can achieve an average velocity of one body length per second. The vehicle is able to carry various sensors or communication equipments, as the payload capacity is about 4 kg. Initial testing shows that the operation time of the buoyancy body is estimated to about 6 hours for free swimming and 90 hours for a pure gliding. The flapping frequency, flapping amplitude, and the number of waves performed across the fin's chord and wave directions can be independently tuned through the proposed control scheme. In general, the present prototype provides a useful platform to study the ray-like swimming motion in a single or combination mode of flapping, undulation and gliding.     

A Three-Dimensional Kinematics Analysis of a Koi Carp Pectoral Fin by Digital Image Processing

Pectoral fins fascinate researchers for their important role in fish maneuvers. By possessing a complicated flexible structure with several fin rays made by a thin film, the fin exhibits a three-dimensional (3D) motion. The complex 3D fin kinematics makes it challenging to study the performance of pectoral fin. Nevertheless, a detailed study on the 3D motion pattern of pectoral fins is necessary to the design and control of a bio-inspired fin rays. Therefore, a highspeed photography system is introduced in this paper to study the 3D motion of a Koi Carp by analyzing the two views of its pectoral fin simultaneously. The key motions of the pectoral fins are first captured in both hovering and retreating. Next, the 3D configuration of the pectoral fins is reconstructed by digital image processing, in which the movement of fin rays during fish retreating and hovering is obtained. Furthermore, the method of Singular Value Decomposition (SVD) is adopted to extract the basic motion patterns of pectoral fins from extensive image sequences, i.e. expansion, bending, cupping, and undulation. It is believed that the movement of the fin rays and the basic patterns of the pectoral fins obtained in the present work can provide a good foundation for the development and control of bionic flexible pectoral fins for underwater propeller.     

Function of the dorsal fin in bluegill sunfish: Motor patterns during four distinct locomotor behaviors

The median fins of fishes are key features of locomotor morphology which function as complex control surfaces during a variety of behaviors. However, very few studies have experimentally assessed median fin function, as most workers focus on axial structures. In particular, the dorsal fin of many teleost fishes possesses both spiny anterior and soft posterior portions which may function separately during locomotion. We analyzed the function of the soft region of the dorsal fin and of the dorsal inclinator (Di) muscles which are the primary muscles responsible for lateral flexion. We used electromyography to measure in vivo Di activity, as well as activity of the red myomeric muscles located at a similar longitudinal position. We quantified motor patterns during four locomotor behaviors: braking and three propulsive behaviors (steady swimming, kick and glide swimming, and C-starts). During the three propulsive swimming behaviors, the timing of Di activity was more similar to that of ipsilateral red myomeric muscle rather than to contralateral myomeric activity, whereas during braking the timing of activity of the Di muscles was similar to that of the contralateral myomeric musculature. During the three propulsive behaviors, when the Di muscles had activity, it was consistent with the function of stiffening the soft dorsal fin to oppose its tendency to bend as a result of the body being swept laterally through the water. In contrast, activity of the Di muscles during braking was consistent with the function of actively flexing the soft dorsal fin towards the side of the fish that had Di activity. Activity of the Di muscles during steady speed swimming was generally sufficient to resist lateral bending of the soft dorsal fin, whereas during high speed kick and glide swimming and C-starts, Di activity was not sufficient to resist the bending caused by resistive forces imposed by the water. Cumulative data from all four behaviors suggest that the Di muscles can be activated independently relative to the myomeric musculature rather than having a single phase relationship with the myomeric muscle common to all of the observed behaviors. © 1996 Wiley-Liss, Inc.     

Locomotion of Gymnarchus Niloticus： Experiment and Kinematics

In addition to forward undulatory swimming, Gymnarchus niloticus can swim via undulations of the dorsal fin while the body axis remains straight; furthermore, it swims forward and backward in a similar way, which indicates that the undulation of the dorsal fin can simultaneously provide bidirectional propulsive and maneuvering forces with the help of the tail fin. A high-resolution Charge-Coupled Device （CCD） imaging camera system is used to record kinematics of steady swimming as well as maneuvering in G. niloticus. Based on experimental data, this paper discusses the kinematics （cruising speed, wave speed, cycle frequency, amplitude, lateral displacement） of forward as well as backward swimming and maneuvering. During forward swimming, the propulsive force is generated mainly by undulations of the dorsal fin while the body axis remains straight. The kinematic parameters （wave speed, wavelength, cycle frequency, amplitude） have statistically significant correlations with cruising speed. In addition, the yaw at the head is minimal during steady swimming. From experimental data, the maximal lateral displacement of head is not more than 1% of the body length, while the maximal lateral displacement of the whole body is not more than 5% of the body length. Another important feature is that G. niloticus swims backwards using an undulatory mechanism that resembles the forward undulatory swimming mechanism. In backward swimming, the increase of lateral displacement of the head is comparatively significant; the amplitude profiles of the propulsive wave along the dorsal fin are significantly different from those in forward swimming. When G. niloticus does fast maneuvering, its body is first bent into either a C shape or an S shape, then it is rapidly unwound in a travelling wave fashion. It rarely maneuvers without the help of the tail fin and body bending.     

中国爬岩鳅属鱼类一新种记述(鲤形目,平鳍鳅科)

记述了采自云南省沾益县德泽乡牛栏江水域的平鳍鳅科爬岩鳅属鱼类1新种,牛栏爬岩鳅 Beaufortia niulanensis Chen,Huang et Yang,sp.nov..其特征:背鳍分支鳍条7,腹鳍分支鳍条21,侧线鳞90～95,胸鳍起点相对于鼻孔和眼前缘之间,背鳍起点相对于腹鳍起点至其基部后缘的中点稍后,肛门位于腹鳍基后缘至臀鳍起点间的中点,腹鳍末端接近肛门,这些特征组合可以将新种与同属的其他2个相近种,即四川爬岩鳅B.szechuanensis(Fang)和中间爬岩鳅B.intermedia Tang et Wang区别开来.     

Parametric Study of an Underwater Finned Propulsor Inspired by Bluespotted Ray

The performance of bluespotted rays was emulated in the design of a bioinspired underwater propulsor in the present work.First,the movement of a live bluespotted ray was captured for the swimming mode and useful information to the biomimetic mechanism design.By virtue of the modular and reeonfigurable design concept,an undulatory fin propulsion prototype was developed.With a proper experimental set-up,orthogonal experiments were conducted to investigate the effect of various fin design parameters on the propulsion speed,thrust,and power of the fish robot.The controllable fin parameters include frequency,amplitude,wavelength,fm shape,and undulatory mode.The significance of these parameters was also determined by using the variance analysis.The results demonstrate that the designed propulsor,imitating bluespotted rays with large expanded undulatory fins,is able to propel itself by changing various kinematic parameters.     

A new species of sexually dimorphic and rheophilic ghost knifefish (Apteronotidae: Gymnotiformes) from the Amazon basin

A new species of ghost knifefish, Apteronotus, is described from high-energy environments in the Rios Mapuera and Trombetas (at Cachoeira Porteira waterfalls), Brazil. X-ray microcomputed tomography (μCT scan) was used to access the internal anatomy of the type series. The new species is distinguished from all congeners by the anteriormost position of the anus, with its posterior margin extending less than one eye diameter beyond the vertical through the caudal limit of the posterior nostril, the low number of anal-fin rays (117–125) and the reduced number of branchiostegal rays (three). A series of modifications associated with secondary sexual dimorphism on the preorbital region of mature males are depicted and discussed. In addition, comments on homologies of the branchiostegal rays in Apteronotidae are provided.     

Functional Morphology of Aquatic Flight in Fishes: Kinematics, Electromyography, and Mechanical Modeling of Labriform Locomotion

Labriform locomotion is the primary swimming mode for many fishesthat use the pectoral fins to generate thrust across a broadrange of speeds. A review of the literature on hydrodynamics,kinematics, and morphology of pectoral fin mechanisms in fishesreveals that we lack several kinds of morphological and kinematicdata that are critical for understanding thrust generation inthis mode, particularly at higher velocities. Several needsinclude detailed three-dimensional kinematic data on speciesthat are pectoral fin swimmers across a broad range of speeds,data on the motor patterns of pectoral fin muscles, and thedevelopment of a mechanical model of pectoral fin functionalmorphology. New data are presented here on pectoral fin locomotionin Gomphosus varius, a labrid fish that uses the pectoral finsat speeds of 1 –6 total body lengths per second. Three-dimensionalkinematic data for the pectoral fins of G. varius show thata typical "drag-based" mechanism is not used in this species.Instead, the thrust mechanics of this fish are dominated bylift forces and acceleration reaction forces. The fin is twistedlike a propeller during the fin stroke, so that angles of attackare variable along the fin length. Electromyographic data onsix fin muscles indicate the sequence of muscle activity thatproduces antagonistic fin abduction and adduction and controlsthe leading edge of the fin. EMG activity in abductors and adductorsis synchronous with the start of abduction and adduction, respectively,so that muscle mechanics actuate the fin with positive work.A mechanical model of the pectoral fin is proposed in whichfin morphometrics and computer simulations allow predictionsof fin kinematics in three dimensions. The transmission of forceand motion to the leading edge of the fin depends on the mechanicaladvantage of fin ray levers. An integrative program of researchis suggested that will synthesize data on morphology, physiology,kinematics, and hydrodynamics to understand the mechanics ofpectoral fin swimming.     

Genetic and thermal effects on the latitudinal variation in the timing of fin development of a fish Oryzias latipes

Heterochrony, an evolutionary change in developmental processes, is one of the major proximate causes of morphological diversity of organisms. It has been reported in the medaka Oryzias latipes that higher-latitude larvae have a genetic tendency to complete fin ray formation at larger body sizes, which results in relatively shorter anal and dorsal fins in adults. However, this latitudinal, heterochronic variation in fin length in the wild may be partially explained by latitudinal differences in thermal environments, if temperatures affect the timing of fin ray formation. Common-environment experiments revealed that the body size at which fin pterygiophore (a basal skeleton of fin rays) formation was completed was larger in higher-latitude larvae than in lower-latitude larvae at all temperatures examined, supporting the proposal that fin ray formation of the former is genetically delayed. However, phenotypic plasticity in response to temperature was also evident; lower temperatures caused delayed fin ray formation until a larger body size had been achieved in both high- and low-latitude larvae. These observations suggest that habitat temperatures also contribute to the latitudinal difference in the timing of fin development, magnifying phenotypic variation in fin length across latitudes. We discuss reasons for this positive covariance between genetic and environmental effects on the latitudinal, heterochronic variation, from the viewpoint of local adaptation and evolution of phenotypic plasticity.     

中国隆头鱼科一新纪录种——雀尖嘴鱼

尖嘴鱼属(Gomphosus)是一群分布于印度洋和太平洋热带珊瑚礁海域鱼类,共有2种,以往在中国海域记录有1种杂色尖嘴鱼(G.varius)。我们在分析20世纪90年代采自中国南海大陆坡的鱼类标本时,发现了该属的另一种雀尖嘴鱼(Gomphosus caeruleus Lacepède,1801),为中国新纪录种。本种的主要鉴别特征为:体呈浅黄褐色(雌)或深黑色(雄);吻部特别延长呈管状;体长为体高的4.2倍,为头长的2.6倍;背鳍Ⅷ-13,臀鳍Ⅲ-11,胸鳍i(不分支)+14(分支);脊椎骨25;鳃盖条7;体被中大圆鳞;侧线完全,在背鳍条的后部下方急剧向下弯折,侧线有孔鳞片27;头部仅鳃盖上部有9枚呈三角状排列的小鳞;背鳍起点前方有鳞8行;背鳍第一至第三鳍棘间的鳍膜具1黑斑;尾鳍截形。     

Re‐evaluation of batoid pectoral morphology reveals novel patterns of diversity among major lineages

Batoids (Chondrichthyes: Batoidea) are a diverse group of cartilaginous fishes which comprise a monophyletic sister lineage to all neoselachians or modern sharks. All species in this group possess anteroposteriorly expanded‐pectoral fins, giving them a unique disc‐like body form. Reliance on pectoral fins for propulsion ranges from minimal (sawfish) to almost complete dependence (skates and rays). A recent study on the diversity of planform pectoral fin shape in batoids compared overall patterns of morphological variation within the group. However, inconsistent pectoral homology prevented the study from accurately representing relationships within and among major batoid taxa. With previous work in mind, we undertook an independent investigation of pectoral form in batoids and evaluated the implications of shape diversity on locomotion and lifestyle, particularly in the skates (Rajoidei) and rays (Myliobatoidei). We used geometric morphometrics with sliding semilandmarks to analyze pectoral fin outlines and also calculate fin aspect ratios (AR), a functional trait linked to locomotion. In agreement with previous work, our results indicated that much of the evolution of batoid pectoral shape has occurred along a morphological axis that is closely related to AR. For species where kinematic data were available, both shape and AR were associated with swimming mode. This work further revealed novel patterns of shape variation among batoids, including strong bimodality of shape in rays, an intermediate location of skate species in the morphospace between benthic/demersal and pelagic rays, and approximately parallel shape trajectories in the benthic/demersal rays and skates. Finally, manipulation of landmarks verified the need for a consistent and accurate definition of homology for the outcome and efficacy of analyses of pectoral form and function in batoids. J. Morphol. 277:482–493, 2016. © 2016 Wiley Periodicals, Inc.     

The locomotory function of the fins in the squid Loligo pealei

Kinematic data of high spatial and temporal resolution, acquired from image sequences of adult long-finned squid, Loligo pealei, during steady swimming in a flume, were used to examine the role of fins and the coordination between fin and jet propulsion in squid locomotion. Fin shape and body outlines were digitized and used to calculate fin wave speed, amplitude, frequency, angle of attack, body deformation, speed, and acceleration. L. pealei were observed to have two fin gait patterns with a transition at 1.4-1.8 mantle lengths per second (Lm s-1) marked by alternation between the two patterns. Fin motion in L. pealei exhibited characteristics of both traveling waves and flapping wings. At low speeds, fin motion was more wave-like; at high speeds, fin motion was more flap-like and was marked by regular periods during which the fins were wrapped tightly against the mantle. Fin cycle frequencies were dependent on swimming speed and gait, and obvious coordination between the fins and jet were observed. Fin wave speed, angle of attack, and body acceleration confirmed the role of fins in thrust production and revealed a role of fins at all swimming speeds by a transition from drag-based to lift-based thrust when fin wave speed dropped below swimming speed. Estimates of peak fin thrust were as high as 0.44-0.96 times peak jet thrust in steady swimming over the range of swimming speeds observed. Fin downstrokes generally contributed more to thrust than did upstrokes, especially at high speeds.     

Ontogenetic variation in fin ray segmentation between latitudinal populations of the medaka, <Emphasis Type="Italic">Oryzias latipes</Emphasis>

Fin rays of ray-finned fishes are composed of multiple bony segments, and each fin ray elongates by adding a new segment to the tip. Therefore, fin ray length is determined by the number of segments and the length of each segment. A comparison of the anal fin rays of a northern and southern wild population of the medaka, Oryzias latipes, revealed that southern fish had more segments per fin ray, resulting in longer anal fins than the northern fish. When fish were reared in a laboratory common environment, segmentation of the fin rays started earlier with respect to body size in the southern fish. In the southern males, moreover, the rate of segment addition accelerated after a certain body size, indicating sexual maturity. These patterns of segment addition during ontogeny were consistent with the patterns of fin ray elongation. Although distal segments tended to be longer, except for the most proximal segment, in both populations, the southern fish had shorter segments than the northern fish at any position on fin rays. These results indicate that the interpopulation variation in fin length is largely due to genetically-based differences in the control of segment addition, and that the length of each segment does not contribute to it. We suspect that fin ray segmentation is regulated by thyroid and sex hormones that differ between populations. We also found that some segments fuse with each other at the base of each fin ray, the functions and mechanisms of which remain unclear.     

A new cave‐dwelling loach,Triplophysa xichouensis sp. nov. (Teleostei Nemacheilidae) from Yunnan,China

A new cave‐dwelling loach of the genus Triplophysa, T. xichouensis, is described from an outlet of a subterranean river in Xisa Town, Xichou County, Yunnan Province, China. It can be distinguished from its congeners by the following characters: dorsal‐fin rays iii, 8; anal‐fin rays ii, 6; pectoral‐fin rays i, 9 or 10; pelvic‐fin rays i, 5 or 6; branched caudal‐fin rays 16(8+8); eyes highly degenerated to a very tiny black dot; dorsal‐fin origin closer to snout tip than to caudal‐fin base and anterior to vertical line of pelvic‐fin origin; pectoral fin length about two‐thirds the distance between pectoral‐fin origin to pelvic‐fin origin; caudal peduncle slender, its length about three times its depth; caudal fin emarginate; body smooth and scaleless; lateral line complete and straight; anterior chamber of air bladder wrapped in dumbbell‐shaped bony capsule and the posterior one well developed, long, oval; intestine short, bending in zigzag shape behind stomach. A key for the cave‐dwelling species of Triplophysa is provided. urn:lsid:zoobank.org:pub:9162FFB1‐7911‐47C3‐AE50‐6A00E9590327     

大麻哈鱼卵黄囊期仔鱼异速生长及其生态学意义

运用实验生态学的方法, 对大麻哈鱼(Oncorhynchus keta Walbaum)卵黄囊期仔鱼的异速生长及器官优先发育在早期生存和环境适应上的生态学意义进行了研究。结果表明, 大麻哈鱼卵黄囊期仔鱼的感觉、摄食, 呼吸和游泳等器官快速分化, 许多关键器官均存在异速生长现象。在身体各部分中, 头部和尾部为正异速生长, 躯干部为负异速生长, 体高有先增大后减小的趋势; 在头部器官中, 眼径、口宽、吻长和眼后头长均为正异速生长; 在游泳器官中, 胸鳍、腹鳍、背鳍、臀鳍、背鳍基、臀鳍基和尾鳍均为正异速生长, 脂鳍为负异速生长, 其中, 腹鳍在全长25.31 mm、12日龄出现生长拐点, 但拐点前后均为正异速生长。大麻哈鱼卵黄囊期仔鱼感觉、摄食, 呼吸和游泳等器官的快速发育, 使出膜后的仔鱼在最短的时间内获得了与早期生存密切相关的各种能力, 对适应复杂多变的外界环境具有重要的生态学意义。