From fish to modern humans--comparative anatomy, homologies and evolution of the head and neck musculature

In a recent paper Diogo (2008) reported the results of the first part of an investigation of the comparative anatomy, homologies and evolution of the head and neck muscles of osteichthyans (bony fish + tetrapods). That report mainly focused on actinopterygian fish, but also compared these fish with certain non-mammalian sarcopterygians. The present paper focuses mainly on sarcopterygians, and particularly on how the head and neck muscles have evolved during the transitions from sarcopterygian fish and non-mammalian tetrapods to monotreme and therian mammals, including modern humans. The data obtained from our dissections of the head and neck muscles of representative members of sarcopterygian fish, amphibians, reptiles, monotremes and therian mammals, such as rodents, tree-shrews, colugos and primates, including modern humans, are compared with the information available in the literature. Our observations and comparisons indicate that the number of mandibular and true branchial muscles (sensu this work) present in modern humans is smaller than that found in mammals such as tree-shrews, rats and monotremes, as well as in reptiles such as lizards. Regarding the pharyngeal musculature, there is an increase in the number of muscles at the time of the evolutionary transition leading to therian mammals, but there was no significant increase during the transition leading to the emergence of higher primates and modern humans. The number of hypobranchial muscles is relatively constant within the therian mammals we examined, although in this case modern humans have more muscles than other mammals. The number of laryngeal and facial muscles in modern humans is greater than that found in most other therian taxa. Interestingly, modern humans possess peculiar laryngeal and facial muscles that are not present in the majority of the other mammalian taxa; this seems to corroborate the crucial role played by vocal communication and by facial expressions in primate and especially in human evolution. It is hoped that by compiling, in one paper, data about the head and neck muscles of a wide range of sarcopterygians, the present work could be useful to comparative anatomists, evolutionary biologists and functional morphologists and to researchers working in other fields such as developmental biology, genetics and/or evolutionary developmental biology.     

On the origin, homologies and evolution of primate facial muscles, with a particular focus on hominoids and a suggested unifying nomenclature for the facial muscles of the Mammalia

The mammalian facial muscles are a subgroup of hyoid muscles (i.e. muscles innervated by cranial nerve VII). They are usually attached to freely movable skin and are responsible for facial expressions. In this study we provide an account of the origin, homologies and evolution of the primate facial muscles, based on dissections of various primate and non‐primate taxa and a review of the literature. We provide data not previously reported, including photographs showing in detail the facial muscles of primates such as gibbons and orangutans. We show that the facial muscles usually present in strepsirhines are basically the same muscles that are present in non‐primate mammals such as tree‐shrews. The exceptions are that strepsirhines often have a muscle that is usually not differentiated in tree‐shrews, the depressor supercilii, and lack two muscles that are usually differentiated in these mammals, the zygomatico‐orbicularis and sphincter colli superficialis. Monkeys such as macaques usually lack two muscles that are often present in strepsirhines, the sphincter colli profundus and mandibulo‐auricularis, but have some muscles that are usually absent as distinct structures in non‐anthropoid primates, e.g. the levator labii superioris alaeque nasi, levator labii superioris, nasalis, depressor septi nasi, depressor anguli oris and depressor labii inferioris. In turn, macaques typically lack a risorius, auricularis anterior and temporoparietalis, which are found in hominoids such as humans, but have muscles that are usually not differentiated in members of some hominoid taxa, e.g. the platysma cervicale (usually not differentiated in orangutans, panins and humans) and auricularis posterior (usually not differentiated in orangutans). Based on our observations, comparisons and review of the literature, we propose a unifying, coherent nomenclature for the facial muscles of the Mammalia as a whole and provide a list of more than 300 synonyms that have been used in the literature to designate the facial muscles of primates and other mammals. A main advantage of this nomenclature is that it combines, and thus creates a bridge between, those names used by human anatomists and the names often employed in the literature dealing with non‐human primates and non‐primate mammals.     

Comparative anatomy, homologies and evolution of the pectoral and forelimb musculature of tetrapods with special attention to extant limbed amphibians and reptiles

The main aim of the present work is to synthesize the information obtained from our dissections of the pectoral and forelimb muscles of representative members of the major extant taxa of limbed amphibians and reptiles and from our review of the literature, in order to provide an account of the comparative anatomy, homologies and evolution of these muscles in the Tetrapoda. The pectoral and forelimb musculature of all these major taxa conform to a general pattern that seems to have been acquired very early in the evolutionary history of tetrapods. Although some muscles are missing in certain taxa, and a clear departure from this general pattern is obviously present in derived groups such as birds, the same overall configuration is easily distinguishable in these taxa. Among the most notable anatomical differences between the groups, one that seems to have relevant evolutionary and functional implications, concerns the distal insertion points of the forearm musculature. In tetrapods, the muscles of the radial and ulnar complexes of the forearm are pleisomorphically mainly inserted onto the radius/ulna or onto the more proximal carpal bones, but in mammals some of these muscles insert more distally onto bones such as the metacarpals. Interestingly, a similar trend towards a more distal insertion of these muscles is also found in some non‐mammalian tetrapod taxa, such as some anurans (e.g. Phyllomedusa). This may be correlated with the acquisition of more subtle digital movement abilities in these latter taxa.     

Taking a look into the orbit of mammalian carnivorans

In this study, we explore the relationship between orbit anatomy and different ecological factors in carnivorous mammals from a phylogenetic perspective. We calculated the frontation (α), convergence (β), and orbitotemporal (Ω) angles of the orbit from 3D coordinates of anatomical landmarks in a wide sample of carnivores with different kinds of visual strategy (i.e. photopic, scotopic, and mesopic), habitat (i.e. open, mixed, and closed), and substrate use (i.e. arboreal, terrestrial, and aquatic). We used Bloomberg's K and Pagel's λ to assess phylogenetic signal in frontation, convergence, and orbitotemporal angles. The association of orbit orientation with skull length and ecology was explored using phylogenetic generalized least squares and phylogenetic manova , respectively. Moreover, we also computed phylomorphospaces from orbit orientation. Our results indicate that there is not a clear association between orbit orientation and the ecology of living carnivorans. We hypothesize that the evolution of the orbit in mammalian carnivores represents a new case of an ecological bottleneck specific to carnivorans. New directions for future research are discussed in light of this new evidence.     

Running,jumping, hunting,and scavenging: Functional analysis of vertebral mobility and backbone properties in carnivorans

Carnivorans are well-known for their exceptional backbone mobility, which enables them to excel in fast running and long jumping, leading to them being among the most successful predators amongst terrestrial mammals. This study presents the first large-scale analysis of mobility throughout the presacral region of the vertebral column in carnivorans. The study covers representatives of 6 families, 24 genera and 34 species. We utilized a previously developed osteometry-based method to calculate available range of motion, quantifying all three directions of intervertebral mobility: sagittal bending (SB), lateral bending (LB), and axial rotation (AR). We observed a strong phylogenetic signal in the structural basis of the vertebral column (vertebral and joint formulae, length proportions of the backbone modules) and an insignificant phylogenetic signal in most characteristics of intervertebral mobility. This indicates that within the existing structure (stabilization of which occurred rather early in different phylogenetic lineages), intervertebral mobility in carnivorans is quite flexible. Our findings reveal that hyenas and canids, which use their jaws to seize prey, are characterized by a noticeably elongated cervical region and significantly higher SB and LB mobility of the cervical joints compared to other carnivorans. In representatives of other carnivoran families, the cervical region is very short, but the flexibility of the neck (both SB and LB) is significantly higher than that of short-necked odd-toed and even-toed ungulates. The lumbar region of the backbone in carnivorans is dorsomobile in the sagittal plane, being on average ~23° more mobile than in artiodactyls and ~38° more mobile than in perissodactyls. However, despite the general dorsomobility, only some representatives of Canidae, Felidae, and Viverridae are superior in lumbar flexibility to the most dorsomobile ungulates. The most dorsomobile artiodactyls are equal or even superior to carnivorans in their ability to engage in dorsal extension during galloping. In contrast, carnivorans are far superior to ungulates in their ability to engage ventral flexion. The cumulative SB in the lumbar region in carnivorans largely depends on the mode of running and hunting. Thus, adaptation to prolonged and enduring pursuit of prey in hyenas is accompanied by markedly reduced SB flexibility in the lumbar region. A more dorsostable run is also a characteristic of the Ursidae, and the peculiar maned wolf. Representatives of Felidae and Canidae have significantly more available SB mobility in the lumbar region. However, they fully engage it only occasionally at key moments of the hunt associated with the direct capture of the prey or when running in a straight line at maximum speed.     

Digastric muscle of the kangaroo: A comparative anatomical study

Although the human digastric muscle is classified as a suprahyoid muscle, none of the digastric muscles in other mammals are classed as suprahyoid in textbooks of veterinary anatomy. The aim of this study was to describe the anatomical relationship of the digastric muscle in a marsupial, the kangaroo, and to consider factors thought to be important in leading to the different position of the muscle in quadrupeds compared with humans. Eight heads of the common wallaroo (Macropus robustus)were used in this study. They were fixed by injection of 10% formalin solution into the carotid arteries and dissected under a stereomicroscope. The digastric muscle in the common wallaroo arose from the paroccipital process of the temporal bone and inserted into the mandible but had no intermediate tendon or any connection with the hyoid bone. It was supplied by both the mandibular and facial nerves. The hyoglossus muscle attached to the inferior surface of the hyoid bone and its ventral border overlapped the mylohyoid muscle. The hypoglossal nerve passed between these two muscles. Therefore, in contrast to humans, the digastric, hyoglossus, and mylohyoid muscles in the kangaroo were all located inferior to the hyoid bone. Differences in head posture and the position of the larynx between kangaroos and humans may account for the observed difference in the digastric muscle's position relative to the hyoid bone between these species. Anat. Rec. 251:346–350, 1998. © 1998 Wiley-Liss, Inc.     

Molecular biological investigations of Brazilian Tunga sp. isolates from man, dogs, cats, pigs and rats

Twenty-four isolates of Tunga were collected from afflicted humans, dogs, cats, pigs and rats in Brazil. To investigate genetic diversity, a hypervariable section of mitochondrial 16S rDNA was amplified using PCR and subsequently sequenced. In order to compare results with another species of the genus Tunga, three isolates of the recently described Tunga trimamillata were also checked. Whereas eleven isolates (five from cats, three from dogs and three from humans) were of identical sequence, thirteen isolates collected from dogs, humans, pigs and rats showed differences in sequence up to 49%, so that the existence of one or more new species of Tunga may be presumed.     

The oblique extraocular muscles in cetaceans: Overall architecture and accessory insertions

The oblique extraocular muscles (EOMs) were dissected in 19 cetacean species and 10 non-cetacean mammalian species. Both superior oblique (SO) and inferior oblique (IO) muscles in cetaceans are well developed in comparison to out-groups and have unique anatomical features likely related to cetacean orbital configurations, swimming mechanics, and visual behaviors. Cetacean oblique muscles originate at skeletal locations typical for mammals: SO, from a common tendinous cone surrounding the optic nerve and from the medially adjacent bone surface at the orbital apex; IO, from the maxilla adjacent to lacrimal and frontal bones. However, because of the unusual orbital geometry in cetaceans, the paths and relations of SO and IO running toward their insertions onto the temporal ocular sclera are more elaborate than in humans and most other mammals. The proximal part of the SO extends from its origin at the apex along the dorsomedial aspect of the orbital contents to a strong fascial connection proximal to the preorbital process of the frontal bone, likely the cetacean homolog of the typical mammalian trochlea. However, the SO does not turn at this connection but continues onward, still a fleshy cylinder, until turning sharply as it passes through the external circular muscle (ECM) and parts of the palpebral belly of the superior rectus muscle. Upon departing this “functional trochlea” the SO forms a primary scleral insertion and multiple accessory insertions (AIs) onto adjacent EOM tendons and fascial structures. The primary SO scleral insertions are broad and muscular in most cetacean species examined, while in the mysticete minke whale (Balaenoptera acutorostrata) and fin whale (Balaenoptera physalus) the muscular SO bellies transition into broad fibrous tendons of insertion. The IO in cetaceans originates from an elongated fleshy attachment oriented laterally on the maxilla and continues laterally as a tubular belly before turning caudally at a sharp bend where it is constrained by the ECM and parts of the inferior rectus which form a functional trochlea as with the SO. The IO continues to a fleshy primary insertion on the temporal sclera but, as with SO, also has multiple AIs onto adjacent rectus tendons and connective tissue. The multiple IO insertions were particularly well developed in pygmy sperm whale (Kogia breviceps), minke whale and fin whale. AIs of both SO and IO muscles onto multiple structures as seen in cetaceans have been described in humans and domesticated mammals. The AIs of oblique EOMs seen in all these groups, as well as the unique “functional trochleae” of cetacean SO and IO seem likely to function in constraining the lines of action at the primary scleral insertions of the oblique muscles. The gimble-like sling formed by SO and IO in cetaceans suggest that the “primary” actions of the cetacean oblique EOMs are not only to produce ocular counter-rotations during up-down pitch movements of the head during swimming but also to rotate the plane containing the functional origins of the rectus muscles during other gaze changes.     

Structural and functional anatomy of the neck musculature of the dog (Canis familiaris)

The morphometric properties and the anatomical relationships of the entire musculature of the canine cervical spine are reported herein. These data were obtained from the dissection of cadavers of six dogs. Total muscle length, muscle weight, fascicle length and angles of pennation were recorded for each muscle comprising the canine cervical spine. Based upon these properties, physiological cross-section area (PCSA) and architectural index were estimated. When scaled by whole body mass, the values of each of these parameters were found to be similar between all dogs. Muscles that course from the cranial neck to the shoulder girdle or the rib cage (e.g. brachiocephalicus and rhomboideus capitis) were found to have relatively long fascicles and low PCSA values and thus appear to be designed for rapid excursions. By contrast, muscles that primarily support the neck and shoulder against gravitational forces (e.g. serratus ventralis and trapezius) were found to have relatively high PCSA values and short fascicle lengths, and thus have the capacity to generate large forces. Differences of morphometry as well as nomenclature were found between the canine and human neck musculature. Nevertheless, many similarities exist; in particular, both species have similar muscles adapted to force generation or large excursions. We thus conclude that the canine neck may be used as a modelling tool for biomechanical investigations of the human cervical region as long as the differences listed are borne in mind.     

A survey of the prevalence and genotypes of Giardia duodenalis infecting household and sheltered dogs

Giardia duodenalis (syn. G. intestinalis, G. lamblia) is an intestinal protozoan parasite that infects humans and a wide range of mammals that includes dogs. Highly significant genetic heterogeneity has been found within this species, while only genotypes from assemblages A and B have zoonotic potential. Although Giardia infection in dogs has been reported worldwide, there is increasing molecular evidence that dogs may be infected with host-specific genotypes (C and D) as well as zoonotic ones (A and B). Therefore, dogs play a role as a potential source of Giardia infection impacting humans and other Canidae. Fecal samples from privately owned dogs and from dogs kept in two shelters in the west-central region of Poland were examined using a microscope and polymerase chain reaction. The total prevalence of Giardia in specimens was 1.9%. Deoxyribonucleic acid was extracted only from two out of the three Giardia-positive fecal samples. After amplification using the G7 and G759 β-giardin primers, 753 bp amplicons were obtained, and both amplification products were sequenced. Sequence and phylogenetic analyses showed that both G. duodenalis isolates were dog-specific genotypes (C and D).     

From fish to modern humans – comparative anatomy, homologies and evolution of the pectoral and forelimb musculature

In a recent study Diogo & Abdala [(2007 ) J Morphol  268 , 504–517] reported the results of the first part of a research project on the comparative anatomy, homologies and evolution of the pectoral muscles of osteichthyans (bony fish and tetrapods). That report mainly focused on actinopterygian fish but also compared these fish with certain non-mammalian sarcopterygians. This study, which reports the second part of the research project, focuses mainly on sarcopterygians and particularly on how the pectoral and forelimb muscles have evolved during the transitions from sarcopterygian fish and non-mammalian tetrapods to monotreme and therian mammals and humans. The data obtained by our own dissections of all the pectoral and forelimb muscles of representative members of groups as diverse as sarcopterygian fish, amphibians, reptiles, monotremes and therian mammals such as rodents, tree-shrews, colugos and primates, including humans, are compared with the information available in the literature. Our observations and comparisons clearly stress that, with regard to the number of pectoral and forelimb muscles, the most striking transition within sarcopterygian evolutionary history was that leading to the origin of tetrapods. Whereas extant sarcopterygian fish have an abductor and adductor of the fin and a largely undifferentiated hypaxial and epaxial musculature, extant salamanders such as Ambystoma have more than 40 pectoral and forelimb muscles. There is no clear increase in the number of pectoral and forelimb muscles within the evolutionary transition that led to the origin of mammals and surely not to that leading to the origin of primates and humans.     

Plantibodies in human and animal health: a review

BackgroundAntibodies are essential part of vertebrates'' adaptive immune system; they can now be produced by transforming plants with antibody-coding genes from mammals/humans. Although plants do not naturally make antibodies, the plant-derived antibodies (plantibodies) have been shown to function in the same way as mammalian antibodies.MethodsPubMed and Google search engines were used to download relevant publications on plantibodies in medical and veterinary fields; the papers were reviewed and findings qualitatively described.ResultsThe process of bioproduction of plantibodies offers several advantages over the conventional method of antibody production in mammalian cells with the cost of antibody production in plants being substantially lesser. Contrary to what is possible with animal-derived antibodies, the process of making plantibodies almost exclusively precludes transfer of pathogens to the end product. Additionally, plants not only produce a relatively high yield of antibodies in a comparatively faster time, they also serve as cost-effective bioreactors to produce antibodies of diverse specificities.ConclusionPlantibodies are safe, cost-effective and offer more advantages over animal-derived antibodies. Methods of producing them are described with a view to inspiring African scientists on the need to embrace and harness this rapidly evolving biotechnology in solving human and animal health challenges on the continent where the climate supports growth of diverse plants.     

Muscular Reconstruction and Functional Morphology of the Forelimb of Early Miocene Sloths (Xenarthra,Folivora) of Patagonia

Early Miocene sloths are represented by a diversity of forms ranging from 38 to 95 kg, being registered mainly from Santacrucian Age deposits in southern‐most shores of Patagonia, Argentina. Their postcranial skeleton differs markedly in shape from those of their closest living relatives (arboreal forms of less than 10 kg), Bradypus and Choloepus. In order to gain insight on functional properties of the Santacrucian sloths forelimb, musculature was reconstructed and a comparative, qualitative morphofunctional analysis was performed, allowing proposing hypotheses about biological role of the limb in substrate preferences, and locomotor strategies. The anatomy of the forelimb of Santacrucian sloths resembles more closely extant anteaters such as Tamandua and Myrmecophaga, due to the robustness of the elements, development of features related to attachment of ligaments and muscles, and conservative, pentadactylous, and strong‐clawed manus. The reconstructed forelimb musculature was very well developed and resembles that of extant Pilosa (especially anteaters), although retaining the basic muscular configuration of generalized mammals. This musculature allowed application of powerful forces, especially in adduction of the forelimb, flexion and extension of the antebrachium, and manual prehension. These functional properties are congruent with both climbing and digging activities, and provide support for proposed Santacrucian sloths as good climbing mammals, possibly arboreal or semiarboreal, being also capable diggers. Their climbing strategies were limited, thus these forms relied mainly on great muscular strength and curved claws of the manus to move cautiously on branches. Anat Rec, 2013. © 2012 Wiley Periodicals, Inc.     

Soft-tissue anatomy of the primates: phylogenetic analyses based on the muscles of the head, neck, pectoral region and upper limb, with notes on the evolution of these muscles

Apart from molecular data, nearly all the evidence used to study primate relationships comes from hard tissues. Here, we provide details of the first parsimony and Bayesian cladistic analyses of the order Primates based exclusively on muscle data. The most parsimonious tree obtained from the cladistic analysis of 166 characters taken from the head, neck, pectoral and upper limb musculature is fully congruent with the most recent evolutionary molecular tree of Primates. That is, this tree recovers not only the relationships among the major groups of primates, i.e. Strepsirrhini {Tarsiiformes [Platyrrhini (Cercopithecidae, Hominoidea)]}, but it also recovers the relationships within each of these inclusive groups. Of the 301 character state changes occurring in this tree, ca. 30% are non-homoplasic evolutionary transitions; within the 220 changes that are unambiguously optimized in the tree, ca. 15% are reversions. The trees obtained by using characters derived from the muscles of the head and neck are more similar to the most recent evolutionary molecular tree than are the trees obtained by using characters derived from the pectoral and upper limb muscles. It was recently argued that since the Pan/Homo split, chimpanzees accumulated more phenotypic adaptations than humans, but our results indicate that modern humans accumulated more muscle character state changes than chimpanzees, and that both these taxa accumulated more changes than gorillas. This overview of the evolution of the primate head, neck, pectoral and upper limb musculature suggests that the only muscle groups for which modern humans have more muscles than most other extant primates are the muscles of the face, larynx and forearm.     

A Model for Transport and Dispersion in the Circulatory System Based on the Vascular Fractal Tree

Materials are distributed throughout the body of mammals by fractal networks of branching tubes. Based on the scaling laws of the fractal structure, the vascular tree is reduced to an equivalent one-dimensional, tube model. A dispersion–convection partial differential equation with constant coefficients describes the heterogeneous concentration profile of an intravascular tracer in the vascular tree. A simple model for the mammalian circulatory system is built in entirely physiological terms consisting of a ring shaped, one-dimensional tube which corresponds to the arterial, venular, and pulmonary trees, successively. The model incorporates the blood flow heterogeneity of the mammalian circulatory system. Model predictions are fitted to published concentration-time data of indocyanine green injected in humans and dogs. Close agreement was found with parameter values within the expected physiological range. © 2003 Biomedical Engineering Society. PAC2003: 8710+e, 8719Hh, 8719Uv     

Prevalence of antibodies against Neospora caninum in dogs from urban areas in Central Poland

Neospora caninum is a protozoan parasite which causes abortion in cattle as well as reproduction problems and neurological disorders in dogs. To assess the prevalence of the parasite in urban dogs in the Mazovian Voivodeship, Central Poland, serum samples from 257 dogs were analyzed for the presence of specific IgG antibodies. The examined dogs visited three private veterinary clinics located in Warsaw due to control tests, vaccinations, or other reasons not directly connected with neosporosis. Using ELISA and Western blot, antibodies against the parasite were detected in 56 out of 257 dogs, giving a prevalence of 21.7%. A greater prevalence was observed in female dogs than in males, 28% and 17.3%, respectively, and the differences were statistically significant (p < 0.05). There were no significant differences in seroprevalence of Neospora infection within the age groups (p > 0.05). This study indicates the presence of N. caninum in the Mazovian Voivodeship, in dogs which live in urban areas and exposure of these dogs to the parasite. The fact that seropositive dogs had no contact with cattle confirms the important role of dogs in the parasite’s epidemiology.