The pectoral fin of Panderichthys and the origin of digits

One of the identifying characteristics of tetrapods (limbed vertebrates) is the presence of fingers and toes. Whereas the proximal part of the tetrapod limb skeleton can easily be homologized with the paired fin skeletons of sarcopterygian (lobe-finned) fish, there has been much debate about the origin of digits. Early hypotheses interpreted digits as derivatives of fin radials, but during the 1990s the idea gained acceptance that digits are evolutionary novelties without direct equivalents in fish fin skeletons. This was partly based on developmental genetic data, but also substantially on the pectoral fin skeleton of the elpistostegid (transitional fish/tetrapod) Panderichthys, which appeared to lack distal digit-like radials. Here we present a CT scan study of an undisturbed pectoral fin of Panderichthys demonstrating that the plate-like 'ulnare' of previous reconstructions is an artefact and that distal radials are in fact present. This distal portion is more tetrapod-like than that found in Tiktaalik and, in combination with new data about fin development in basal actinopterygians, sharks and lungfish, makes a strong case for fingers not being a novelty of tetrapods but derived from pre-existing distal radials present in all sarcopterygian fish.     

An autopodial-like pattern of Hox expression in the fins of a basal actinopterygian fish

Comparative analyses of Hox gene expression and regulation in teleost fish and tetrapods support the long-entrenched notion that the distal region of tetrapod limbs, containing the wrist, ankle and digits, is an evolutionary novelty. Data from fossils support the notion that the unique features of tetrapod limbs were assembled over evolutionary time in the paired fins of fish. The challenge in linking developmental and palaeontological approaches has been that developmental data for fins and limbs compare only highly derived teleosts and tetrapods; what is lacking are data from extant taxa that retain greater portions of the fin skeletal morphology considered primitive to all bony fish. Here, we report on the expression and function of genes implicated in the origin of the autopod in a basal actinopterygian, Polyodon spathula. Polyodon exhibits a late-phase, inverted collinear expression of 5' HoxD genes, a pattern of expression long considered a developmental hallmark of the autopod and shown in tetrapods to be controlled by a 'digit enhancer' region. These data show that aspects of the development of the autopod are primitive to tetrapods and that the origin of digits entailed the redeployment of ancient patterns of gene activity.     

A Devonian tetrapod-like fish and the evolution of the tetrapod body plan

The relationship of limbed vertebrates (tetrapods) to lobe-finned fish (sarcopterygians) is well established, but the origin of major tetrapod features has remained obscure for lack of fossils that document the sequence of evolutionary changes. Here we report the discovery of a well-preserved species of fossil sarcopterygian fish from the Late Devonian of Arctic Canada that represents an intermediate between fish with fins and tetrapods with limbs, and provides unique insights into how and in what order important tetrapod characters arose. Although the body scales, fin rays, lower jaw and palate are comparable to those in more primitive sarcopterygians, the new species also has a shortened skull roof, a modified ear region, a mobile neck, a functional wrist joint, and other features that presage tetrapod conditions. The morphological features and geological setting of this new animal are suggestive of life in shallow-water, marginal and subaerial habitats.     

Three-dimensional limb joint mobility in the early tetrapod Ichthyostega

The origin of tetrapods and the transition from swimming to walking was a pivotal step in the evolution and diversification of terrestrial vertebrates. During this time, modifications of the limbs—particularly the specialization of joints and the structures that guide their motions—fundamentally changed the ways in which early tetrapods could move. Nonetheless, little is known about the functional consequences of limb anatomy in early tetrapods and how that anatomy influenced locomotion capabilities at this very critical stage in vertebrate evolution. Here we present a three-dimensional reconstruction of the iconic Devonian tetrapod Ichthyostega and a quantitative and comparative analysis of limb mobility in this early tetrapod. We show that Ichthyostega could not have employed typical tetrapod locomotory behaviours, such as lateral sequence walking. In particular, it lacked the necessary rotary motions in its limbs to push the body off the ground and move the limbs in an alternating sequence. Given that long-axis rotation was present in the fins of tetrapodomorph fishes, it seems that either early tetrapods evolved through an initial stage of restricted shoulder and hip joint mobility or that Ichthyostega was unique in this respect. We conclude that early tetrapods with the skeletal morphology and limb mobility of Ichthyostega were unlikely to have made some of the recently described Middle Devonian trackways.     

Evidence that mechanisms of fin development evolved in the midline of early vertebrates

The origin of paired appendages was a major evolutionary innovation for vertebrates, marking the first step towards fin- (and later limb-) driven locomotion. The earliest vertebrate fossils lack paired fins but have well-developed median fins, suggesting that the mechanisms of fin development were assembled first in the midline. Here we show that shark median fin development involves the same genetic programs that operate in paired appendages. Using molecular markers for different cell types, we show that median fins arise predominantly from somitic (paraxial) mesoderm, whereas paired appendages develop from lateral plate mesoderm. Expression of Hoxd and Tbx18 genes, which specify paired limb positions, also delineates the positions of median fins. Proximodistal development of median fins occurs beneath an apical ectodermal ridge, the structure that controls outgrowth of paired appendages. Each median fin bud then acquires an anteroposteriorly-nested pattern of Hoxd expression similar to that which establishes skeletal polarity in limbs. Thus, despite their different embryonic origins, paired and median fins utilize a common suite of developmental mechanisms. We extended our analysis to lampreys, which diverged from the lineage leading to gnathostomes before the origin of paired appendages, and show that their median fins also develop from somites and express orthologous Hox and Tbx genes. Together these results suggest that the molecular mechanisms for fin development originated in somitic mesoderm of early vertebrates, and that the origin of paired appendages was associated with re-deployment of these mechanisms to lateral plate mesoderm.     

An exceptional Devonian fish from Australia sheds light on tetrapod origins

The transition from fishes to tetrapods was one of the most dramatic events in the evolution of vertebrates, but many pivotal fossils are incomplete, resulting in gaps in the data that are used for phylogenetic reconstruction. Here we present new observations from the most complete, acid-prepared Devonian tetrapodomorph fish yet discovered, Gogonasus, which was previously placed just crownward of Kenichthys and rhizodontids, the most primitive taxa on the tetrapod lineage. Unexpectedly, Gogonasus shows a mosaic of plesiomorphic and derived tetrapod-like features. Whereas the braincase and dermal cranial skeleton exhibit generalized morphologies with respect to Eusthenopteron or Panderichthys, taxa that are traditionally considered to be phyletically close to tetrapods, the presence of a deeply invaginated, wide spiracle, advanced internal spiracular architecture and near-horizontal hyomandibula are specialized features that are absent from Eusthenopteron. Furthermore, the pectoral fin skeleton of Gogonasus shares several features with that of Tiktaalik, the most tetrapod-like fish. A new phylogenetic analysis places Gogonasus crownward of Eusthenopteron as the sister taxon to the Elpistostegalia. Aspects of the basic tetrapod limb skeleton and middle ear architecture can now be traced further back within the tetrapodomorph radiation.     

The pelvic fin and girdle of Panderichthys and the origin of tetrapod locomotion

One of the most marked transformations in the vertebrate transition to land was that of fins to limbs. This transformation involved not only the generation of morphological novelties (digits, sacrum) but also a shift in locomotory dominance from the pectoral to the pelvic appendage. Despite its importance, the transformation from pelvic fin to hindlimb is the least studied and least well-documented part of this transformation, which is bracketed by the osteolepiform Eusthenopteron and the early tetrapods Ichthyostega and Acanthostega, but is not directly illuminated by any intermediate form. Panderichthys is the closest tetrapod relative currently represented by complete fossils, but its pelvic fin skeleton has not been described. Here, I present the only known articulated pelvic fin endoskeleton and associated partial pelvis of Panderichthys. The pelvic girdle is even less tetrapod-like than that of the osteolepiform Eusthenopteron, but the pelvic fin endoskeleton shares derived characteristics with basal tetrapods despite being more primitive than the pectoral fin of Panderichthys. The evolution of tetrapod locomotion appears to have passed through a stage of body-flexion propulsion, in which the pelvic fins played a relatively minor anchoring part, before the emergence of hindlimb-powered propulsion in the interval between Panderichthys and Acanthostega.     

The cranial endoskeleton of Tiktaalik roseae

Among the morphological changes that occurred during the 'fish-to-tetrapod' transition was a marked reorganization of the cranial endoskeleton. Details of this transition, including the sequence of character acquisition, have not been evident from the fossil record. Here we describe the braincase, palatoquadrate and branchial skeleton of Tiktaalik roseae, the Late Devonian sarcopterygian fish most closely related to tetrapods. Although retaining a primitive configuration in many respects, the cranial endoskeleton of T. roseae shares derived features with tetrapods such as a large basal articulation and a flat, horizontally oriented entopterygoid. Other features in T. roseae, like the short, straight hyomandibula, show morphology intermediate between the condition observed in more primitive fish and that observed in tetrapods. The combination of characters in T. roseae helps to resolve the relative timing of modifications in the cranial endoskeleton. The sequence of modifications suggests changes in head mobility and intracranial kinesis that have ramifications for the origin of vertebrate terrestriality.     

Ventastega curonica and the origin of tetrapod morphology

The gap in our understanding of the evolutionary transition from fish to tetrapod is beginning to close thanks to the discovery of new intermediate forms such as Tiktaalik roseae. Here we narrow it further by presenting the skull, exceptionally preserved braincase, shoulder girdle and partial pelvis of Ventastega curonica from the Late Devonian of Latvia, a transitional intermediate form between the 'elpistostegids' Panderichthys and Tiktaalik and the Devonian tetrapods (limbed vertebrates) Acanthostega and Ichthyostega. Ventastega is the most primitive Devonian tetrapod represented by extensive remains, and casts light on a part of the phylogeny otherwise only represented by fragmentary taxa: it illuminates the origin of principal tetrapod structures and the extent of morphological diversity among the transitional forms.     

Evolutionary origins of vertebrate appendicular muscle

The evolution of terrestrial tetrapod species heralded a transition in locomotor strategies. While most fish species use the undulating contractions of the axial musculature to generate propulsive force, tetrapods also rely on the appendicular muscles of the limbs to generate movement. Despite the fossil record generating an understanding of the way in which the appendicular skeleton has evolved to provide the scaffold for tetrapod limb musculature, there is, by contrast, almost no information as to how this musculature arose. Here we examine fin muscle formation within two extant classes of fish. We find that in the teleost, zebrafish, fin muscles arise from migratory mesenchymal precursor cells that possess molecular and morphogenetic identity with the limb muscle precursors of tetrapod species. Chondrichthyan dogfish embryos, however, use the primitive mechanism of direct epithelial somitic extensions to derive the muscles of the fin. We conclude that the genetic mechanism controlling formation of tetrapod limb muscles evolved before the Sarcopterygian radiation.     

在辐射和对流条件下肋片传热的研究

本文研究了在辐射和对流条件下梯形肋片的传热,应用四阶龙格-库塔法和牛顿-拉伐森迭代法进行数值求解,得出肋片沿肋高方向的温度分布;进一步分析了导热系数、辐射率、肋根温度、肋高和肋间距等参数对肋片传热的影响。比较了考虑辐射矩形肋、三角形肋和不考虑幅射的梯形肋的传热情况。结果对工程设计具有重要意义。     

大鳍鳠臀鳍和脂鳍的比较组织学研究

大鳍鳠臀鳍和脂鳍均是皮肤特化的产物．臀鳍的组织结构可分为表皮层、胶原纤维层、胶原下层和鳍条４部分；脂鳍可分为表皮层、胶原纤维层和脂肪层．臀鳍和脂鳍的表层均与皮肤的表皮相似，而发达的胶原纤维层与皮肤真皮的致密层相当．臀鳍的胶原下层、脂鳍的脂肪层与皮下层相当，而鳍条则是矿化的结缔组织．从特化程度来看，脂鳍的特化程度不如臀鳍高，表明脂鳍应是奇鳍褶的痕迹器官．     

基于单刃-圆锥面刀具的三维整体外翅片成形机理

采用单刃-圆锥面刀具加工三维整体外翅片管,研究了翅片成形过程及圆锥面半径对翅片成形的影响,分析了翅片微观形貌的形成机理及加工参数对翅片形貌的影响.翅片成形机理在于单刃-圆锥面刀具切削时能将工件表层金属与基体分离但并不去除,被分离的金属在圆锥面的挤压下形成翅片.翅片成形过程可分为犁切分离、滑动挤压和成翅3个阶段,较大的圆弧半径有利于翅片的成形.翅片顶部有一条贯穿整个翅片的微裂缝,使翅片在微观上成为双翅片;随着犁切深度的增加,翅片厚度增加;随着进给量的增大,翅片间距增加.     

空气外掠圆孔翅片管的流动与换热数值模拟

以矩形平翅片作为比较对象,采用数值模拟方法研究了空气外掠三对称大直径圆孔翅片表面的流动与传热性能,获得了不同Re(雷诺)数时矩形平翅片和三对称大直径圆孔翅片表面的速度场、温度场和Nu(努塞尔)数分布.平翅片的模拟结果与实验数据的最大误差小于10%,证明了该模拟方法的正确性.研究结果表明:当气流Re=1 610～6 440时,三对称大直径圆孔翅片的表面传热系数比平翅片提高25%以上.证明该圆孔翅片是一种适用于翅片管式制冷换热器且传热效果优越的片型.     

Mechanical design and implementation of a new biomimetic robot fish

A mechanical design method of robot fish is introduced in this paper.Based on this method an antonomous 3-Dimension(3D)locomotion robot fish with two pectoral fins and a caudal fin is developed.The pectoral fin mechanism has 3 degrees of freedom(3-DOFs),which enables the robot fish to realize yawing and pitching motions by controlling two pectoral fins.And the caudal fin mechanism is designed based on fish body wave curve fitting.The forward velocity can be adjusted by changing the caudal mechanism's oscillating frequency.Finally a physical implementation of the robot fish and experimental results are given.     

计算肋片散热的一种改进方法

通过对二维肋片导热机理的分析，提出了一种改进的计算二维肋片散热和温度分布的一维近似方法。这一方法与经典的一维近似方法一样简单实用，并可套用经典方法的所有公式和图表，分别用改进的一维近似方法、经典一维方法和二维方法对工程中几种常见肋片进行了计算并加以比较。结果表明，一维近似方法较经典一维方法的精度有明显提高。     

Tetrapod-like middle ear architecture in a Devonian fish

Few fossils show the incipient stages of complex morphological transformations. For example, the earliest stages in the remodelling of the spiracular tract and suspensorium (jaw suspension) of osteolepiform fishes into the middle ear of tetrapods have remained elusive. The most primitive known tetrapods show a middle ear architecture that is very different from osteolepiforms such as Eusthenopteron, with little indication of how this transformation took place. Here we present an analysis of tetrapod middle ear origins that is based on a detailed study of Panderichthys, the immediate sister taxon of tetrapods. We show that the spiracular region is radically transformed from osteolepiforms and represents the earliest stages in the origin of the tetrapod middle ear architecture. The posterior palatoquadrate of Panderichthys is completely tetrapod-like and defines a similarly tetrapod-like spiracular tract. The hyomandibula has lost its distal portion, representing a previously unrecognized advance towards a stapes-like morphology. This spiracular specialization suggests that the middle ear of early tetrapods evolved initially as part of a spiracular breathing apparatus.     

三种大管径翅管式换热器传热与阻力特性的试验研究

为考察不同翅片形式以及管排数对空气侧强化传热的影响，分别对9排和12排带平直、开缝、纵向涡3种翅片形式共6个翅片管换热器元件空气侧的传热及阻力性能进行了试验研究，发现开缝翅片管换热器的传热性能高于带纵向涡翅片管换热器和平直翅片管换热器，但阻力相应也增加，带纵向涡发生器的翅片传热性能略低于开缝翅片，但阻力却只比平直翅片稍大，这表明采用这种翅片形式可获得较好的综合性能．不同管排数的同一片型换热器的努塞尔数及阻力因子相差很小，可以认为与管排数无关．在试验的雷诺数范围内针对各个试件整理出了传热和阻力的经验关联式，可为相关的理论研究和工程应用提供参考．     

管壳式冷凝器内低螺纹管的凝结传热模型

以蒸气在垂直壁面和水平圆管上的凝结传热特性为基础，建立了蒸汽在基管，翅片及翅顶上的凝结传热模型，对影响低螺纹管传热的结构参数翅片间距，翅片夹角以及当量翅高进行了计算分析，进而建立了低螺纹管管簇的凝结传热模型，得出了低螺纹管管簇的平均冷凝传热系数与管排数和管列数之间的关系式。