Ventilatory mechanics from maniraptoran theropods to extant birds

Shared behavioural, morphological and physiological characteristics are indicative of the evolution of extant birds from nonavian maniraptoran dinosaurs. One such shared character is the presence of uncinate processes and respiratory structures in extant birds. Recent research has suggested a respiratory role for these processes found in oviraptorid and dromaeosaurid dinosaurs. By measuring the geometry of fossil rib cage morphology, we demonstrate that the mechanical advantage, conferred by uncinate processes, for movements of the ribs in the oviraptorid theropod dinosaur, Citipati osmolskae, basal avialan species Zhongjianornis yangi, Confuciusornis sanctus and the more derived ornithurine Yixianornis grabaui, is of the same magnitude as found in extant birds. These skeletal characteristics provide further evidence of a flow-through respiratory system in nonavian theropod dinosaurs and basal avialans, and indicate that uncinate processes are a key adaptation facilitating the ventilation of a lung air sac system that diverged earlier than extant birds.     

The asymmetry of the carpal joint and the evolution of wing folding in maniraptoran theropod dinosaurs

In extant birds, the hand is permanently abducted towards the ulna, and the wrist joint can bend extensively in this direction to fold the wing when not in use. Anatomically, this asymmetric mobility of the wrist results from the wedge-like shape of one carpal bone, the radiale, and from the well-developed convexity of the trochlea at the proximal end of the carpometacarpus. Among the theropod precursors of birds, a strongly convex trochlea is characteristic of Coelurosauria, a clade including the highly derived Maniraptora in addition to tyrannosaurs and compsognathids. The shape of the radiale can be quantified using a ‘radiale angle’ between the proximal and distal articular surfaces. Measurement of the radiale angle and reconstruction of ancestral states using squared-change parsimony shows that the angle was small (15°) in primitive coelurosaurs but considerably larger (25°) in primitive maniraptorans, indicating that the radiale was more wedge-shaped and the carpal joint more asymmetric. The radiale angle progressively increased still further within Maniraptora, with concurrent elongation of the forelimb feathers and the forelimb itself. Carpal asymmetry would have permitted avian-like folding of the forelimb in order to protect the plumage, an early advantage of the flexible, asymmetric wrist inherited by birds.     

Cardio‐pulmonary anatomy in theropod dinosaurs: Implications from extant archosaurs

Although crocodilian lung and cardiovascular organs are markedly less specialized than the avian heart and lung air‐sac system, all living archosaurs possess four‐chambered hearts and heterogeneously vascularized, faveolar lungs. In birds, normal lung function requires extensive, dorsally situated nonvascularized abdominal air‐sacs ventilated by an expansive sternum and specially hinged costal ribs. The thin walled and voluminous abdominal air‐sacs are supported laterally and caudally to prevent inward (paradoxical) collapse during generation of negative (inhalatory) pressure: the synsacrum, posteriorly directed, laterally open pubes and specialized femoral‐thigh complex provide requisite support and largely prevent inhalatory collapse. In comparison, theropod dinosaurs probably lacked similarly enlarged abdominal air‐sacs, and skeleto‐muscular modifications consistent with their ventilation. In the absence of enlarged, functional abdominal air‐sacs, theropods were unlikely to have possessed a specialized bird‐like, air‐sac lung. The likely absence of bird‐like pulmonary function in theropods is inconsistent with suggestions of cardiovascular anatomy more sophisticated than that of modern crocodilians. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Convergent dental adaptations in the serrations of hypercarnivorous synapsids and dinosaurs

Theropod dinosaurs are well known for having a ziphodont dentition: serrated, blade-shaped teeth that they used for cutting through prey. Serrations along the carinae of theropod teeth are composed of true denticles, a complex arrangement of dentine, enamel, and interdental folds. This structure would have supported individual denticles and dissipated the stresses associated with feeding. These particular serrations were previously thought to be unique to theropod dinosaurs and some other archosaurs. Here, we identify the same denticles and interdental folds forming the cutting edges in the teeth of a Permian gorgonopsian synapsid, extending the temporal and phylogenetic distribution of this dental morphology. This remarkable instance of convergence not only represents the earliest record of this adaptation to hypercarnivory but also demonstrates that the first iteration of this feature appeared in non-mammalian synapsids. Comparisons of tooth serrations in gorgonopsians with those of earlier synapsids and hypercarnivorous mammals reveal some gorgonopsians acquired a complex tissue arrangement that differed from other synapsids.     

Growth patterns in brooding dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian condition

The timing of sexual maturation in non-avian dinosaurs is not known. In extant squamates and crocodilians it occurs in conjunction with the initial slowing of growth rates as adult size is approached. In birds (living dinosaurs) on the other hand, reproductive activity begins well after somatic maturity. Here we used growth line counts and spacing in all of the known brooding non-avian dinosaurs to determine the stages of development when they perished. It was revealed that sexual maturation occurred well before full adult size was reached-the primitive reptilian condition. In this sense, the life history and physiology of non-avian dinosaurs was not like that of modern birds. Palaeobiological ramifications of these findings include the potential to deduce reproductive lifespan, fecundity and reproductive population sizes in non-avian dinosaurs, as well as aid in the identification of secondary sexual characteristics.     

早期鸟类及其近亲的腹肋腔:大小和功能(英文)

对中生代鸟类及与其亲缘关系密切的非鸟兽脚类腹肋腔的大小和形态进行了研究。据报道,在兽脚类中,腹肋腔的大小与个体大小正相关。然而与早期报道相反,基干鸟类的腹肋腔按个体大小比例似乎比非鸟兽脚类的大,这表明在缺乏前后向充分发育的胸骨时,腹肋可能作为一种补偿性结构,具有附着肌肉的功能。像鳄鱼一样,具有胸骨的早期鸟类的近端一排腹肋与胸骨相关联,表明这两种结构在功能上可能是一个整体;但在所有腹肋结构中都没有发现任何用来附着肌肉的痕迹,所以这一观点还没有此方面证据的支持。或者,大型的腹肋腔可能与呼吸功能以及鸟类飞翔需要的大量氧气供应有关。由于这些细弱骨骼结构的保存往往不完整,好的样本太少,以至于无法从对腹肋资料的统计分析中得出重要的结论。     

Taxonomic identification bias does not drive patterns of abundance and diversity in theropod dinosaurs

The ability of palaeontologists to correctly diagnose and classify new fossil species from incomplete morphological data is fundamental to our understanding of evolution. Different parts of the vertebrate skeleton have different likelihoods of fossil preservation and varying amounts of taxonomic information, which could bias our interpretations of fossil material. Substantial previous research has focused on the diversity and macroevolution of non-avian theropod dinosaurs. Theropods provide a rich dataset for analysis of the interactions between taxonomic diagnosability and fossil preservation. We use specimen data and formal taxonomic diagnoses to create a new metric, the Likelihood of Diagnosis, which quantifies the diagnostic likelihood of fossil species in relation to bone preservation potential. We use this to assess whether a taxonomic identification bias impacts the non-avian theropod fossil record. We find that the patterns of differential species abundance and clade diversity are not a consequence of their relative diagnosability. Although there are other factors that bias the theropod fossil record that are not investigated here, our results suggest that patterns of relative abundance and diversity for theropods might be more representative of Mesozoic ecology than often considered.     

Evolution of olfaction in non-avian theropod dinosaurs and birds

Little is known about the olfactory capabilities of extinct basal (non-neornithine) birds or the evolutionary changes in olfaction that occurred from non-avian theropods through modern birds. Although modern birds are known to have diverse olfactory capabilities, olfaction is generally considered to have declined during avian evolution as visual and vestibular sensory enhancements occurred in association with flight. To test the hypothesis that olfaction diminished through avian evolution, we assessed relative olfactory bulb size, here used as a neuroanatomical proxy for olfactory capabilities, in 157 species of non-avian theropods, fossil birds and living birds. We show that relative olfactory bulb size increased during non-avian maniraptoriform evolution, remained stable across the non-avian theropod/bird transition, and increased during basal bird and early neornithine evolution. From early neornithines through a major part of neornithine evolution, the relative size of the olfactory bulbs remained stable before decreasing in derived neoavian clades. Our results show that, rather than decreasing, the importance of olfaction actually increased during early bird evolution, representing a previously unrecognized sensory enhancement. The relatively larger olfactory bulbs of earliest neornithines, compared with those of basal birds, may have endowed neornithines with improved olfaction for more effective foraging or navigation skills, which in turn may have been a factor allowing them to survive the end-Cretaceous mass extinction.     

Basal dinosauriform and theropod dinosaurs from the mid–late Norian (Late Triassic) of Poland: implications for Triassic dinosaur evolution and distribution

The rise of dinosaurs during the Triassic is a widely studied evolutionary radiation, but there are still many unanswered questions about early dinosaur evolution and biogeography that are hampered by an unevenly sampled Late Triassic fossil record. Although very common in western North America and parts of South America, dinosaur (and more basal dinosauriform) remains are relatively rare in the Upper Triassic deposits of Europe, making any new discoveries critically important. One of the most diverse dinosauriform assemblages from Europe comes from the Por?ba site in Poland, a recently described locality with exposures of the Zb?szynek Beds, which have a palynomorph assemblage characteristic for the mid–late Norian in the biostratigraphic schemes of the Germanic Basin. Using a synapomorphy‐based approach, we evaluate several isolated dinosauriform specimens from Por?ba. This assemblage includes a silesaurid, a herrerasaurid and remains of another type of theropod (potentially a neotheropod). The Por?ba herrerasaurid is the first record of this rare group of primitive dinosaurs from Europe and one of the youngest records worldwide, whereas the silesaurid is the youngest record of a silesaurid from Europe. These findings indicate that silesaurids persisted alongside true dinosaurs into the mid–late Norian of Europe and that silesaurid–herrerasaurid–neotheropod assemblages (which are also known from the Norian of North America, at low latitudes) were more widespread geographically and latitudinally than previously thought. Silesaurid–herrerasaurid–neotheropod assemblages may have been a common ecological structuring of dinosaurs during their early evolution, and their widespread distribution may indicate weak palaeolatitudinal controls on early dinosaur biogeography during the latest Triassic.     

The structural mechanics and evolution of aquaflying birds

Mass‐specific bone strength was examined in the forelimb and hindlimb of 64 species of birds to determine if aquaflying birds (which utilize the wings for propulsion underwater) differ in their skeletal strength compared with other avian taxa. Long bone strengths were estimated from cross‐sectional measurements. Compared with the expectation from geometric similarity, humeral section modulus in volant birds scales nearly isometrically, while femoral strength scales with significant positive allometry. Penguin mass‐specific humeral strength is greatly elevated, but the average humeral strength in species that are propelled by the wings in both air and water do not differ from the values calculated in non‐diving taxa. However, amphibious flyers have gracile femora. Comparative analyses using independent contrasts were utilized to examine the impact of phylogenetic signal. The residual measured for the penguin–procellariiform humeral strength contrast was larger in magnitude (residual of 2.14) than at any other node in the phylogeny. The data strongly indicate that the transition from an amphibious flight condition to a fully aquatic condition involves greater changes in mechanical factors than the transition from purely aerial locomotion to amphibious wing use. There remains the possibility that a historical effect, such as ancestral body size, has impacted the mechanical scaling of penguins. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 687–698.     

Aspects of comparative cranial mechanics in the theropod dinosaurs Coelophysis, Allosaurus and Tyrannosaurus

The engineering analysis technique finite element analysis (FEA) is used here to investigate cranial stress and strain during biting and feeding in three phylogenetically disparate theropod taxa: Coelophysis bauri , Allosaurus fragilis and Tyrannosaurus rex . Stress patterns are generally similar in all taxa with the ventral region of the skull tensed whilst the dorsal aspect is compressed, although the skull is not purely behaving as a cantilever beam as there is no discernible neutral region of bending. Despite similarities, stress patterns are not wholly comparable: there are key differences in how certain regions of the skull contain stress, and it is possible to link such differences to cranial morphology. In particular, nasal morphology can be explained by the stress patterns revealed here. Tyrannosaurus models shear and compress mainly in the nasal region, in keeping with the indistinguishably fused and expanded morphology of the nasal bones. Conversely Allosaurus and Coelophysis models experience peak shear and compression in the fronto-parietal region (which is tightly interdigitated and thickened in the case of Allosaurus ) yet in contrast the nasal region is lightly stressed, corresponding to relatively gracile nasals and a frequently patent internasal suture evident in Allosaurus . Such differences represent alternate mechanical specializations between taxa that may be controlled by functional, phylogenetic or mechanical constraints. Creation of finite element models placed in a phylogenetic context permits the investigation of the role of such mechanical character complexes in the cranium of nonavian theropods and the lineage leading towards modern birds.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 309–316.     

Rib cage mechanics during quiet breathing and exercise in humans

Kenyon, C. M., S. J. Cala, S. Yan, A. Aliverti, G. Scano, R. Duranti, A. Pedotti, and Peter T. Macklem. Rib cage mechanics during quiet breathing and exercise in humans. J. Appl. Physiol. 83(4): 1242-1255, 1997.Duringexercise, large pleural, abdominal, and transdiaphragmatic pressureswings might produce substantial rib cage (RC) distortions. We used athree-compartment chest wall model (J. Appl.Physiol. 72: 1338-1347, 1992) to measuredistortions of lung- and diaphragm-apposed RC compartments (RCp andRCa) along with pleural and abdominal pressures in five normal men. RCpand RCa volumes were calculated from three-dimensional locations of 86 markers on the chest wall, and the undistorted (relaxation) RCconfiguration was measured. Compliances of RCp and RCa measured duringphrenic stimulation against a closed airway were 20 and 0%,respectively, of their values during relaxation. There was marked RCdistortion. Thus nonuniform distribution of pressures distorts the RCand markedly stiffens it. However, during steady-state ergometerexercise at 0, 30, 50, and 70% of maximum workload, RC distortionswere small because of a coordinated action of respiratory muscles, sothat net pressures acting on RCp and RCa were nearly the samethroughout the respiratory cycle. This maximizes RC compliance andminimizes the work of RC displacement. During quiet breathing, plots ofRCa volume vs. abdominal pressure were to the right of the relaxationcurve, indicating an expiratory action on RCa. We attribute this topassive stretching of abdominal muscles, which more thancounterbalances the insertional component of transdiaphragmatic pressure.

     

Evolutionary changes in pubic orientation in dinosaurs are more strongly correlated with the ventilation system than with herbivory

Among dinosaurs, the pubis has convergently retroverted four times in Maniraptora (Theropoda) and once in Ornithischia. Although a clear correlation has not been demonstrated, it has been previously proposed that two traits were related to pubic retroversion: the reduced importance of cuirassal ventilation, and a herbivorous diet. Here, we analyse the possible influence of these traits on pubis orientation. Cuirassal ventilation was plesiomorphically present as an accessory ventilation mechanism in Dinosauria and was powered by the M. ischiotruncus, which was probably connected to a propubic pelvis. Cuirassal ventilation was reduced in both Ornithischia and Maniraptora, some of which also evolved herbivory. According to our results, cuirassal ventilation is more strongly correlated with pubic orientation than herbivory. The retroversion of the pubis during the evolution of birds resulted in major changes in the musculature of the tail. These changes increased the efficiency of the pubocaudalis muscles, which enhanced the birds’ capability for take‐off from the ground. The release of the evolutionary constraint on pubic orientation through changes in the ventilatory system can therefore be considered to be an important step in bird evolution.     

Challenges in teaching the mechanics of breathing to medical and graduate students

The mechanics of breathing has always been a difficult topic for some medical and graduate students. The subject is very quantitative and contains a number of concepts that some students have trouble with, including physical principles such as pressure, flow, volume, resistance, elasticity, and compliance. Apparently, present-day students find the subject more difficult than students of 20 years ago. A possible reason for this is that the teaching of elementary physics in high school and college is now given less emphasis, whereas other topics, such as molecular biology, receive a great deal of attention. Another factor may be that many of us grew up building radios and other such devices, whereas modern students tend to plug in an electronic unit with little idea of its function. Some examples of misconceptions of present-day students who have taken our course are given. To help the weaker students, we now include a primer at the beginning of our handout for the course that covers simple physical principles. Examples of some of the most difficult concepts for students are given.     

欧亚类禽猪流感病毒病原学研究进展

欧亚类禽猪流感病毒被认为是引起下次流感大流行可能性最大的动物流感病毒之一。1979年,第一次在猪群中分离到欧亚类禽猪流感病毒,该病毒随后在欧洲流行。欧亚类禽猪流感病毒从2005年起在我国猪群中广泛存在,偶尔可突破种属屏障感染人;在江苏、河北和湖南等地区已出现人感染欧亚类禽猪流感病毒,由此对公共卫生和生猪养殖业均有一定的威胁和影响。本文综述了欧亚类禽猪流感病毒的流行情况、病原学特征及其相关致病分子机制研究进展,旨在为欧亚类禽猪流感病毒的风险评估提供科学依据。     

Changes in mechanics of breathing during experimental cough and sneeze in anaesthetized cats

Pleural pressure, airflow and tidal volume during experimental cough and sneeze elicited by mechanical stimulation of the tracheobronchial and nasal mucous membranes were investigated in fifty anaesthetized cats (pentobarbital, 40 mg/kg i.p.). Pressure-volume, pressure-flow and flow-volume relations were studied during these expulsive processes. In comparison to quiet breathing there was a decrease in dynamic lung compliance in both respiratory tract reflexes (p less than 0.001), especially in their expiratory phases. As compared to quiet breathing, the total work of breathing was significantly increased (p less than 0.001) in cough (20 times) as well as in sneeze (13 times). The total lung resistance increased markedly (p less than 0.001) in both cough and sneeze compared to quiet breathing. In these expulsive processes there was also a high "cough index" (resistance calculated from the peak flow and instantaneous pressure). The flow-volume curve in cough, in contradistinction to sneeze, indicated a significantly reduced airflow of the end of expiration (at 85% of the expired volume), demonstrating a concomitant bronchoconstriction.