Elbow Extensor Muscles in Humans and Chimpanzees: Adaptations to Different Uses of the Upper Extremity in Hominoid Primates.

The anatomical and functional characteristics of the elbow extensor muscles (triceps brachii and anconeus) have not been widely studied in non-human hominoid primates, despite their great functional importance. In the present study, we have analyzed the muscle architecture and the expression of the myosin heavy chain (MHC) isoforms in the elbow extensors in humans and chimpanzees. Our main objective was to identify differences in these muscles that could be related to the different uses of the upper extremity in the two species. In five humans and five chimpanzees, we have analyzed muscle mass (MM), muscle fascicle length (MFL), and the physiological cross-sectional area (PCSA). In addition, we have assessed the expression of the MHC isoforms by RT-PCR. We have found high MM and PCSA values and higher expression of the MHC-IIx isoform in the triceps brachii of chimpanzees, while in humans, the triceps brachii has high MFL values and a higher expression of the MHC-I and MHC-IIa isoforms. In contrast, there were no significant differences between humans and chimpanzees in any of the values for the anconeus. These findings could be related to the participation of the triceps brachii in the locomotion of chimpanzees and to the use of the upper extremity in manipulative functions in humans. The results obtained in the anconeus support its primary function as a stabilizer of the elbow joint in the two species.

mRNA expression of myosin heavy chain isoforms in the sphenomandibularis portion of the temporalis muscle / Expresión de ARNm de las isoformas de la cadena pesada de la miosina en la porción esfenomandibular del músculo temporal

SUMMARY: The main objective of this study was to analyze by real-time quantitative polymerase chain reaction (RT-qPCR) the expression patterns of the myosin heavy chain (MHC) isoforms (MHC-I, MHC-IIa, MHC-IIx) in the sphenomandibularis portion of the temporalis muscle. We expected to find differences between the sphenomandibularis and the other portions of the temporalis that could be related to the functional characteristics of the sphenomandibularis identified by electromyography. We dissected the right temporalis muscle of ten adult human individuals (five men and five women). Samples of the anterior and posterior temporalis and of the sphenomandibularis portion were obtained from each dissected muscle. These samples were analyzed by RT-qPCR to determine the percentages of expression of the MHC-I, MHC-IIa and MHC-IIx isoforms. No significant differences were identified between the anterior and the posterior temporalis in the expression patterns of the MHC-I, MHC-IIa and MHC-IIx isoforms. However, there were significant differences between the sphenomandibularis and the anterior temporalis. Specifically, the sphenomandibularis portion had a higher percentage of expression of the MHC-I isoform (P=0.04) and a lower percentage of expression of the MHC-IIx isoform (P=0.003). The pattern of expression that we observed in the sphenomandibularis reflects a greater resistance to fatigue, a lower contraction speed, and a lower capacity of force generation in the sphenomandibularis compared to the anterior temporalis. These characteristics are consistent with electromyographic findings on the functional differences between these two portions.

RESUMEN: El principal objetivo de este estudio fue analizar mediante real-time quantitative polymerase chain reaction (RT-qPCR) los patrones de expresión de las isoformas de la cadena pesada de la miosina (MHC-I, MHC-IIa y MHC-IIx) en la porción esfenomandibular del músculo temporal. Se esperó encontrar diferencias entre el esfenomandibular y las otras porciones del músculo temporal que se pudieran relacionar con las características funcionales del esfenomandibular, identificadas mediante electromiografía. Para obtener estos resultados, se diseccionó el músculo temporal derecho en diez humanos adultos (cinco hombres y cinco mujeres) y se obtuvieron muestras de la porción anterior y posterior del músculo temporal y de su porción esfenomandibular. Estas muestras fueron analizadas mediante RT-qPCR para determinar los porcentajes de expresión de las isoformas MHC-I, MHC- IIa y MHC-IIx. No se identificaron diferencias significativas de los patrones de expresión entre la porción anterior y la porción posterior del músculo temporal, pero sí que se observaron diferencias significativas entre la porción anterior del músculo temporal y su porción esfenomandibular. Concretamente, la porción esfenomandibular presentó un mayor porcentaje de expresión de la isoforma MHC-I (P=0.04) y un menor porcentaje de expresión de la isoforma MHC-IIx (P=0.003). El patrón de expresión que hemos observado en la porción esfenomandibular del músculo temporal refleja una mayor resistencia a la fatiga, una velocidad de contracción más lenta y una menor capacidad de generar fuerza si se compara esta porción con la porción anterior del músclo temporal. Estas características son consistentes con las diferencias funcionales que presentan estas dos porciones, que han sido descritas mediante electromiografía.

Quantitative anatomical analysis of the carpal tunnel in women and men.

PURPOSE: The main objective of this study was to identify anatomical differences between men and women in the absolute and relative size of the carpal tunnel (CT), its inner structures, and related external anatomical structures in order to shed light on the higher prevalence of CT syndrome (CTS) in women. BASIC PROCEDURES: We have dissected the forearms and hands of ten men and ten women and compared the cross-sectional area (CSA) of the CT between the two sexes. The size of the CT relative to the tendons passing through it, the median nerve (MN), and to hand and wrist size was also compared between men and women. MAIN FINDINGS: The absolute CSAs of the CT and other parameters were larger in men than in women. The CSA of the CT relative to the length of the capitate bone was also larger in men. However, no significant differences were observed between men and women in the size of the CT relative to its inner structures. PRINCIPAL CONCLUSIONS: The size of the inner structures of the CT are in proportion to that of the CT itself in both sexes. These findings suggest that the etiology of CTS seems to be primarily related to workload and personal traits.

Anatomical and molecular analyses of the deltoid muscle in chimpanzees (Pan troglodytes) and modern humans (Homo sapiens): Similarities and differences due to the uses of the upper extremity.

In the deltoid muscles of Pan troglodytes and Homo sapiens, we have analyzed the muscle architecture and the expression of the myosin heavy chain (MHC) isoforms. Our aim was to identify differences between the two species that could be related to their different uses of the upper limb. The deltoid muscle of six adult Pan troglodytes and six adult Homo sapiens were dissected. The muscle fascicle length (MFL) and the physiological cross-sectional area (PCSA) of each muscle were calculated in absolute and normalized values. The expression pattern of the MHC-I, MHC-IIa and MHC-IIx isoforms was analyzed in the same muscles by real-time polymerase chain reaction. Only the acromial deltoid (AD) presented significant architectural differences between the two species, with higher MFL values in humans and higher PCSA values in chimpanzees. No significant differences in the expression pattern of the MHC isoforms were identified. The higher PCSA values in the AD of Pan troglodytes indicate a greater capacity of force generation in chimpanzees than in humans, which may be related to a greater use of the upper limb in locomotion, specifically in arboreal locomotion like vertical climbing. The functional differences between chimpanzees and humans in the deltoid muscle are more related to muscle architecture than to a differential expression of MHC isoforms.

3D geometric morphometric analysis of the distal radius insertion sites of the palmar radiocarpal ligaments indicates a relationship between wrist anatomy and unique locomotor behavior in hylobatids.

OBJECTIVES: The objective of this study is to explore the anatomical differences in the insertion sites of the palmar radiocarpal ligaments between hylobatids and other hominoids that may be related to their different locomotor behaviors. MATERIALS AND METHODS: The morphology of the insertion sites of the palmar radiocarpal ligaments was analyzed with three-dimensional geometric morphometrics (3D GM) in the distal radial epiphysis of 44 hylobatids, 25 Pan, 31 Gorilla and 15 Pongo. RESULTS: Relative to other hominoids, hylobatid insertion sites of the palmar radiocarpal ligaments were relatively larger and the insertion site of the short radiolunate ligament had a palmar orientation. DISCUSSION: Larger palmar radiocarpal ligaments in hylobatids can help stabilize the wrist during the radial and ulnar displacement that occurs in ricochetal brachiation, the characteristic locomotor behavior of hylobatids, and compensate for the large traction loads on the wrist during extended-elbow vertical climbing.

The Tongue in Three Species of Lemurs: Flower and Nectar Feeding Adaptations.

The mobility of the primate tongue allows for the manipulation of food, but, in addition, houses both general sensory afferents and special sensory end organs. Taste buds can be found across the tongue, but the ones found within the fungiform papillae on the anterior two thirds of the tongue are the first gustatory structures to come into contact with food, and are critical in making food ingestion decisions. Comparative studies of both the macro and micro anatomy in primates are sparse and incomplete, yet there is evidence that gustatory adaptation exists in several primate taxa. One is the distally feathered tongues observed in non-destructive nectar feeders, such as Eulemur rubriventer. We compare both the macro and micro anatomy of three lemurid species who died of natural causes in captivity. We included the following two non-destructive nectar feeders: Varecia variegata and Eulemur macaco, and the following destructive flower feeder: Lemur catta. Strepsirrhines and tarsiers are unique among primates, because they possess a sublingua, which is an anatomical structure that is located below the tongue. We include a microanatomical description of both the tongue and sublingua, which were accomplished using hematoxylin-eosin and Masson trichrome stains, and scanning electron microscopy. We found differences in the size, shape, and distribution of fungiform papillae, and differences in the morphology of conical papillae surrounding the circumvallate ones in all three species. Most notably, large distinct papillae were present at the tip of the tongue in nectar-feeding species. In addition, histological images of the ventro-apical portion of the tongue displayed that it houses an encapsulated structure, but only in Lemur catta case such structure presents cartilage inside. The presence of an encapsulated structure, coupled with the shared morphological traits associated with the sublingua and the tongue tip in Varecia variegata and Eulemur macaco, point to possible feeding adaptations that facilitate non-destructive flower feeding in these two lemurids.

Expression of myosin heavy chain isoform mRNA transcripts in the masseter and medial pterygoid muscles / Expresión de transcripciones de ARNm de isoforma de cadena pesada de miosina en los músculos masetero y pterigoideo medial

SUMMARY: Both the masseter and medial pterygoid muscles elevate the mandible, raising the lower jaw by acting simultaneously on the lateral and medial surfaces of the mandibular ramus. Nevertheless, electromyographic studies indicate that these muscles, as well as the superficial and deep heads of the masseter, act in a different way during mastication. We have analyzed by real time quantitative polymerase chain reaction (RT-qPCR) the expression of myosin heavy chain (MHC) isoforms in the masseter and medial pterygoid muscles in humans in order to identify possible differences in the expression patterns that may be related to functional differences identified with electromyography. Our findings indicate that the expression pattern of MHC isoforms in the two muscles is characteristic of fast and powerful phasic muscles. We have also observed a high percentage of expression of the MHC-IIx isoform and the expression of the MHC-M isoform at the mRNA level in both muscles, an isoform that does not translate into protein in the masticatory muscles of humans. The high percentage of expression of the MHC-IIx isoform in humans can be related to a high contractile speed of the masseter and medial pterygoid in humans. On the other hand, the low percentage of expression of the MHC-M isoform at the mRNA level in both muscles can be related to the complex evolutionary process that has reduced the size and force of the masticatory muscles in humans.

RESUMEN: Los músculos masetero y pterigoideo medial elevan la mandíbula actuando de forma simultánea sobre las caras lateral y medial de su rama. Sin embargo, los estudios electromiográficos indican que estos dos músculos actúan de forma diferente durante la masticación, de la misma forma que lo hacen las porciones superficial y profunda del músculo masetero. En el presente estudio hemos analizado mediante PCR en tiempo real la expresión de las isoformas de la cadena pesada de la miosina o myosin heavy chain (MHC) en los músculos masetero y pterigoideo medial en humanos, con la finalidad de identificar diferencias en los patrones de expresión que se puedan relacionar con las diferencias funcionales identificadas con la electromiografía. Nuestros resultados indican que el patrón de expresión de las isoformas de la MHC en los dos músculos es la característica de los músculos rápidos y potentes. También hemos observado un elevado porcentaje de expresión de la isoforma MHC-IIx y la expresión a nivel de ARNm de la isoforma MHC-M en los dos músculos, una isoforma que no se detecta a nivel de proteína en los músculos masticadores humanos. El elevado porcentaje de expresión de la isoforma MHC-IIx que hemos observado se puede relacionar con una elevada velocidad de contracción de los músculos masetero y pterigoideo medial en los humanos. Por otro lado, el bajo porcentaje de expresión de la isoforma MHC-M a nivel de ARNm en ambos músculos se puede relacionar con los procesos evolutivos complejos que han reducido el tamaño y la fuerza de los músculos masticadores en los humanos.

Effects of Captivity on the Morphology of the Insertion Sites of the Palmar Radiocarpal Ligaments in Hominoid Primates.

The environmental conditions of captive hominoid primates can lead to modifications in several aspects of their behavior, including locomotion, which can then alter the morphological characteristics of certain anatomical regions, such as the knee or wrist. We have performed tridimensional geometric morphometrics (3D GM) analyses of the distal radial epiphysis in wild and captive gorillas, chimpanzees, and orangutans. Our objective was to study the morphology of the insertion sites of the palmar radiocarpal ligaments, since the anatomical characteristics of these insertion sites are closely related to the different types of locomotion of these hominoid primates. We have identified significant differences between the wild and captive specimens that are likely related to their different types of locomotion. Our results indicate that the habitat conditions of captive hominoid primates may cause them to modify their locomotor behavior, leading to a greater use of certain movements in captivity than in the wild and resulting in the anatomical changes we have observed. We suggest that creating more natural environments in zoological facilities could reduce the impact of these differences and also increase the well-being of primates raised in captive environments.

Quantitative shape analysis of the deltoid tuberosity of modern humans (Homo sapiens) and common chimpanzees (Pan troglodytes).

PURPOSE: To identify anatomical differences in the deltoid tuberosity of Homo sapiens and Pan troglodytes, potentially relating to the different uses of the forelimb in these two phylogenetically related species. BASIC PROCEDURES: We have used three-dimensional geometric morphometrics (3D GM) to analyze the deltoid tuberosity of scanned humeri from 30 H. sapiens and 27 P. troglodytes. We also used the 3D scans of the humeri to calculate the surface area of the deltoid tuberosity. Finally, we dissected the deltoid muscles of three H. sapiens and three P. troglodytes to determine the relative mass and the physiological cross-sectional area (PCSA) of each part of the muscle. MAIN FINDINGS: The 3D GM analysis of the deltoid tuberosity identified an anteroposterior enlargement of the P. troglodytes tuberosity, with a lateral displacement of the middle segment, whereas in H. sapiens, there was a distal displacement of the middle segment. Muscle architecture analysis indicated higher normalized values ​​of the PCSA of the clavicular and acromial deltoid in P. troglodytes. PRINCIPAL CONCLUSIONS: The anatomical features observed in our P. troglodytes specimens serve to strengthen the three parts of the deltoid muscle. This fact can be related to the use of the forelimb in locomotion, both arboreal and knuckle-walking, in this species. Humans use the forelimb mainly in manipulative tasks, so they do not develop - as do chimpanzees - the anatomical features that increase the deltoid force. Our findings have shown that the different uses of the forelimb in modern humans and common chimpanzees can affect both muscle architecture and bone morphology, either jointly or separately.

Expression of myosin heavy chain isoforms mRNA transcripts in the temporalis muscle of common chimpanzees (Pan troglodytes).

PURPOSE: The common chimpanzee (Pan troglodytes) is the primate that is phylogenetically most closely related to humans (Homo sapiens). In order to shed light on the anatomy and function of the temporalis muscle in the chimpanzee, we have analyzed the expression patterns of the mRNA transcripts of the myosin heavy chain (MyHC) isoforms in different parts of the muscle. BASIC PROCEDURES: We dissected the superficial, deep and sphenomandibularis portions of the temporalis muscle in five adult P. troglodytes and quantified the expression of the mRNA transcripts of the MyHC isoforms in each portion using real-time quantitative polymerase chain reaction. MAIN FINDINGS: We observed significant differences in the patterns of expression of the mRNA transcripts of the MyHC-IIM isoform between the sphenomandibularis portion and the anterior superficial temporalis (33.6% vs 47.0%; P=0.032) and between the sphenomandibularis portion and the anterior deep temporalis (33.6% vs 43.0; P=0.016). We also observed non-significant differences between the patterns of expression in the anterior and posterior superficial temporalis. PRINCIPAL CONCLUSIONS: The differential expression patterns of the mRNA transcripts of the MyHC isoforms in the temporalis muscle in P. troglodytes may be related to the functional differences that have been observed in electromyographic studies in other species of primates. Our findings can be applicable to the fields of comparative anatomy, evolutionary anatomy, and anthropology.

Expression of MyHC isoforms mRNA transcripts in different regions of the masseter and medial pterygoid muscles in chimpanzees.

OBJECTIVE: The aim of this study is to examine the expression pattern of the different myosin heavy chain (MyHC) isoforms in the masseter and medial pterygoid muscles by real time quantitative polymerase chain reaction (RT-qPCR) to obtain information at molecular level which can be related to the functional characteristics of these two muscles. DESIGN: The masseter, deep and superficial portion, and medial pterygoid muscles of five adult Pan troglodytes were dissected in order to obtain samples of the anterior and posterior regions of each portion of the masseter and of the medial pterygoid. The expression of MyHC isoforms mRNA transcripts was analyzed by RT-qPCR. RESULTS: No significant differences in expression of MyHC isoforms between the masseter and the medial pterygoid were found. In contrast, when comparing the superficial and the deep portion of the masseter, we found that the MyHC-IIM isoform was expressed at a significantly higher level in the superficial portion. CONCLUSIONS: The superficial portion of the masseter and the medial pterygoid muscle have the same expression pattern regarding the different MyHC isoforms. On the other hand, the deep portion of the masseter, which is activated mainly during lateral and repositioning movements of the mandible, has a lower MyHC-IIM isoform expression than the superficial portion. Our findings provide new data on functional aspects of the masseter and medial pterygoid that can complement results obtained by other techniques.

Quantification of Myosin Heavy Chain Isoform mRNA Transcripts in the Supraspinatus Muscle of Vertical Clinger Primates.

Vertical clinging is a specialized form of locomotion characteristic of the primate family Callitrichidae. Vertical clinging requires these pronograde primates to maintain a vertical posture, so the protraction of their forelimbs must resist gravity. Since pronograde primates usually move as horizontal quadrupeds, we hypothesized that the supraspinatus muscle of vertical clingers would present specific characteristics related to the functional requirements imposed on the shoulder area by vertical clinging. To test this hypothesis, we quantified by real-time quantitative polymerase chain reaction the mRNA transcripts of myosin heavy chain (MHC) isoforms in the supraspinatus muscle of 15 species of pronograde primates, including vertical clingers. Our results indicate that the supraspinatus of vertical clingers has a specific expression pattern of the MHC isoforms, with a low expression of the transcripts of the slow MHC-I isoform and a high expression of the transcripts of the fast MHC-II isoforms. We conclude that these differences can be related to the particular functional characteristics of the shoulder in vertical clingers, but also to other anatomical adaptations of these primates, such as their small body size.

Like father, like son: assessment of the morphological affinities of A.L. 288-1 (A. afarensis), Sts 7 (A. africanus) and Omo 119-73-2718 (Australopithecus sp.) through a three-dimensional shape analysis of the shoulder joint.

The postcranial evidence for the Australopithecus genus indicates that australopiths were able bipeds; however, the morphology of the forelimbs and particularly that of the shoulder girdle suggests that they were partially adapted to an arboreal lifestyle. The nature of such arboreal adaptations is still unclear, as are the kind of arboreal behaviors in which australopiths might have engaged. In this study we analyzed the shape of the shoulder joint (proximal humerus and glenoid cavity of the scapula) of three australopith specimens: A.L. 288-1 (A. afarensis), Sts 7 (A. africanus) and Omo 119-73-2718 (Australopithecus sp.) with three-dimensional geometric morphometrics. The morphology of the specimens was compared with that of a wide array of living anthropoid taxa and some additional fossil hominins (the Homo erectus specimen KNM-WT 15000 and the H. neanderthalensis specimen Tabun 1). Our results indicate that A.L. 288-1 shows mosaic traits resembling H. sapiens and Pongo, whereas the Sts 7 shoulder is most similar to the arboreal apes and does not present affinities with H. sapiens. Omo 119-73-2718 exhibits morphological affinities with the more arboreal and partially suspensory New World monkey Lagothrix. The shoulder of the australopith specimens thus shows a combination of primitive and derived traits (humeral globularity, enhancement of internal and external rotation of the joint), related to use of the arm in overhead positions. The genus Homo specimens show overall affinities with H. sapiens at the shoulder, indicating full correspondence of these hominin shoulders with the modern human morphotype.

Shape analysis of the proximal humerus in orthograde and semi-orthograde primates: correlates of suspensory behavior.

"Suspensory locomotion" is an expression that encompasses a series of specialized forms of locomotion that mainly orthograde primates use to achieve below-branch traveling. It implies a number of features in the entire body associated with the use of the forelimb in overhead positions. The glenohumeral joint is one of the main joints involved in effective suspensory locomotion, being subject to a delicate balance between the high degree of mobility and stabilization needed to successfully engage in suspensory behaviors. Here, we present a 3D geometric morphometric study that explores the form of the proximal humerus of six orthograde and semi-orthograde genera (Hylobates, Pongo, Pan, Gorilla, Ateles, and Lagothrix) and a pronograde genus, Colobus, to determine to what extent suspensory locomotor requirements are driving the shape of this epiphysis. Results show the presence of a morphocline related to degree of suspension in the shape of the articular surface, with highly suspensory taxa (i.e., Hylobates) exhibiting particular morphological traits at the articular surface that provide a greater range of circumduction. The placement and orientation of the rotator cuff muscles' insertion sites on the tubercles appear associated with the divergent forces operating at the joint in quadrupedal or above-head use of the hand.

Brief communication: Developmental versus functional three-dimensional geometric morphometric-based modularity of the human proximal humerus.

The proximal humerus is formed by three secondary ossification centers during the postnatal trajectory of the human infant. The ossification centers later grow into the structures of the articular surface, major tubercle, and minor tubercle. There is a purported functional division between the articular surface and the tubercles, with the articular surface mainly responsible for the range of movement of the shoulder joint, and the tubercles bearing the insertions of the rotator cuff muscles, mainly devoted to securing the joint against humeral displacement. Using three-dimensional geometric morphometrics, we tested the presence of such developmental and functional divisions in the proximal humerus, applying the RV coefficient of Escoufier to these a priori hypothesized modules. Our results indicate that the proximal humerus might be a generally integrated structure. However, a weak signal for modular configuration was present, with slightly stronger support for the two modules depicting the boundaries between the purported functional regions of the epiphysis: the articular surface and the tubercles.

Brief communication: Morphological effects of captivity: A geometric morphometric analysis of the dorsal side of the scapula in captive-bred and wild-caught Hominoidea.

Many osteological collections from museums and research institutions consist mainly of remains from captive-bred animals. The restrictions related to the space of their enclosures and the nature of its substrate are likely to affect the locomotor and postural behaviors of captive-bred animals, which are widely considered uninformative regarding bone morphology and anatomical adaptations of wild animals, especially so in the case of extant great apes. We made a landmark-based geometric morphometrics analysis of the dorsal side of the scapular bone of both wild-caught and captive-bred great apes to clarify the effect of captivity on the morphology of a bone greatly involved in locomotion. The comparison suggested that captivity did not have a significant effect on the landmark configuration used, neither on average scapular shape nor shape variability, being impossible to distinguish the scapulae of a captive-bred animal from that of a wild-caught one. This indicates that the analyzed scapulae from captive Hominoidea specimens may be used in morphological or taxonomic analyses since they show no atypical morphological traits caused by living conditions in captivity.

3D geometric morphometric analysis of the proximal epiphysis of the hominoid humerus.

In this study we perform a three-dimensional geometric morphometric (3D GM) analysis of the proximal epiphysis of the humerus in extant great apes, including humans, in order to accurately describe the functional anatomical differences between these taxa. In addition, a fossil hominin specimen of Australopithecus afarensis was included in a multivariate GM analysis in order to test the potential of this methodological approach for making locomotor inferences from fossil remains. The results obtained show significant differences in proximal humeral morphology among the taxa studied, which had thus far largely remained unnoticed. Based on morphofunctional considerations, these anatomical differences can be correlated to differences in the locomotor repertoires of the taxa, thus confirming that the proximal humerus is suitable for constructing paleobiological inferences about locomotion. Modern humans display markedly divergent features, which set them apart from both the extant great apes and the fossil hominin A. afarensis. The morphology of the proximal epiphysis of the humerus of the latter more closely resembles that of the orangutans, thus suggesting that despite hindlimb adaptations to bipedalism, the forelimb of this taxon was still functionally involved in arboreal behaviors, such as climbing or suspension.