Upright posture of the human and the morphologic evolution of the musculi extensores digitorum pedis with reference to evolutionary myology

A brief literature review is made of the morphological changes in the bones of the lower limbs of man, which are the result of his upright walk. The author's task has been to study the morphological changes of Mm. extensores digitorum pedis from the viewpoint of evolutionary myology. The following material has been studied: Lower limbs of adults, 151 less than or equal to N less than or equal to 358. Pelvic limbs of Marsupialia, Insectivora, and non-hominide primates; N = 122. Lower limbs of human embryos and fetuses; N = 71. The following acknowledgements are the author's own studies. They begin with an evolutionary-myological study of m. extensor hallucis longus and of m. extensor digitorum longus, together with m. peroneus tertius.     

Upright posture of man and morphologic evolution of the musculi extensores digitorum pedis with reference to evolutionary myology. III

The following anatomical objects were studied with regard to myology during evolution: M. extensor hallucis longus (MEHL), M. extensor digitorum longus (MEDL) with M. peroneus tertius (MP III), M. peroneus brevis (MPB) with M. peroneus digiti V (MPD V), M. extensor hallucis brevis (MEHB), M. extensor digitorum brevis (MEDB), and the Retinaculum musculorum extensorum imum (RMEI). The study was carried out by the preparation of 3 different groups of material. The 1st group consists of lower extremities of humans. The number of the extremities differs for the particular objects between 151 and 358 (see page 381). The 2nd group of material consists of 122 Membra pelvina from Marsupialia, Insectivora, and Primates. Table 1 shows as well the mammalian species as the number of the studied extremities. The extremities of the 1st and 2nd group were preserved in an manner suitable for a macroscopic preparation. The 3rd group of material consists of 71 lower extremities from embryos and fetus. The lower legs and feet were stained either according to the method described by Morel and Bassal with eosin added or according to Weigert. From this material, complete series of cross sections were prepared. Table 2 shows the age of the embryos (VCL [mm]) as well as the number of the studied extremities. It is important that up to the age of 46 mm VCL the difference in the age of the embryos usually amounts from 0.5 to 1.0 mm. This small difference in the age of the embryos and fetus allows a very good follow up of the changes in construction during the organogenesis. The comparison of the 3 different groups shows the following changes for the above mentioned muscles: The M. extensor hallucis longus (MEHL) is a muscle which is not split. The same result applies for its tendon which inserts at the distal phalanx of the hallux. This primitive form of the muscle amounts actually to 51.12% in human beings. In 48.88% of the cases, additional tendons and muscles are formed by the MEHL. Most of these supplements are positioned on the medial side of the main tendon, only a few lie to the lateral side. For the supplement tendons, the medial one as well as the lateral one occasionally possess a muscle belly. The muscle of the medial tendon is split off from the proximal margin of the MEHL. The muscle of the lateral tendon is split off from the distal margin of the MEHL.(ABSTRACT TRUNCATED AT 400 WORDS)     

Morphologic evolution of the musculi extensores digitorum and abductor pollicis longus in man. Part I: Introduction, methodology, M. extensor digitorum

The author discusses first of all the theoretical principles of the evolutionary myology and gives a detailed explanation on the 3 types of studies included in this complex method. The investigated material is distributed into 3 groups, which correspond to the above mentioned 3 types of studies: 1. The variations of the muscles, object of research, are studied on 200 upper limbs of adults and cn 100 limbs of human fetuses. The crown to heel length of the latter and the number of the studied limbs could be seen on the Table on page 823. 2. The comparative-anatomic research is performed on 122 limbs of animals. The different species of animals as well as the number of the examined limbs are given on the table on page 824. 3. The organogenetic material includes 18 upper limbs of human embryos and fetuses, the age of which could be seen on the table on page 825. After expressing his gratitute to all, who have assisted him in his work the author gives the already known literature data related to the phylogenesis and ontogenesis of the muscles studied by him. His own investigations start with searching of m. extensor digitorum in man, as special attention is paid both to the muscles' bellies and their tendons and the connexus intertendinei.     

A Darwinian evolutionary system. II. Experiments on protein evolution and evolutionary aspects of the genetic code

On the basis of the principles of Darwinian evolutionary systems laid out earlier, a system is constructed which simulates protein evolution. Two types of situations are studied: adaptation to highest possible alkalinity (“alkalinity model”), and adaptation to an arbitrary sequence (“sequence model”). No restrictions in adaptability were found for the (comparably special) alkalinity model, but severe restrictions were found for the sequence model. Approximately 15% of all possible evolutionary paths from one amino acid to another turned out to be impossible, in the sense that no chain of intermediate steps exists which leads to a higher fitness level, in this case an increased chemical similarity of the two amino acids.The evolutionary efficiency of the natural genetic code was also investigated by comparing it with two classes of artificially constructed codes: semi-random and random codes. It was found that the natural code possesses the highest evolutionary efficiency, given by the mean number of generations required to reach identity in 5 of 10 sites, if originally all 10 were different. Closest to the natural code in evolutionary efficiency were the random codes, next, the semi-random codes.This pattern could be explained by a theoretical measure, called the code efficiency. The most important component of the code efficiency is the percentage of impossible paths. The natural code is far superior to the other code types in this respect. However, the random codes are superior to the natural code with respect to the mean shortest path length of the possible paths, the other important component of the code efficiency.It is suggested that the natural genetic code might have arisen from a semi-random code during a process of optimizing several of its features, of which the evolutionary efficiency is a very important one; or that the natural code is the most efficient edition of a large variety of semi-random codes which originated by chance.     

Tongue evolution in the lungless salamanders,family plethodontidae. II. Function and evolutionary diversity

A recently presented model of tongue projection dynamics is used to generate a series of predictions concerning morphologies to be expected under selection for increased distance of projection, increased speed of projection, and increased directional versatility. A general understanding of biomechanical events and the model are used as points of departure for making specific predictions concerning details of structure in skeletal, muscular and connective tissue components of the tongue and associated structures. Comparative methods are used to examine these predictions in the genera of plethodontid salamanders. These salamanders are known to project their tongues to different degrees, and this knowledge is used to test the hypotheses concerning morphological specialization. Three distinct groups of plethodontid salamanders have evolved specializations for long distance projection, and these genera differ from one another in important ways in respect to specific character complexes. For example, the tropical genera and Hydromantes use CBII as the major force transmission element in the skeleton, while Eurycea and its allies use CBI in this role. Hydromantes differs from both in having a uniquely proportioned and structured hyobranchial skeleton and associated musculature. Less extreme specializations for tongue projection are found in different combinations in three other groups. Finally, two distinct groups of generalized species having only limited tongue projection capabilities are recognized, each having a unique complex of inter-related features. Each of these eight groups is recognized and characterized as a functional mode, and hypotheses concerning the biomechanical meaning of the character complexes of each are formulated.     

The parasitic ecology of the rodent mite Myobia musculi. II. Genetic factors.

Mice of 4 strains, C57BL/6J, CD2F1(BALB/cCum female times DBA/2Cum male), BRVR/SrCr and BSVS/SrCr, were monospecifically infested with an equal number of Myobia musculi. A single skin field (7mm diam) in the dorsal midline slightly posterior to the eyes was used for periodic observation of mite populations. Mite counts were made by direct microscopic examination of the skin field at 10-da intervals. Data were obtained on mice thus infested and observed for 130 da. Between-strain comparisons differed significantly in the number of Myobia maintained in all cases except between BRVR and BSVS strains. Factors bearing on heritable differences in susceptibility were discussed.     

The human extensor digitorum profundus muscle with comments on the evolution of the primate hand

Forelimb dissections on 14 genera of anthropoids including humans and 17 cases of human aneuploids has revealed a high incidence of “atavistic” musculature (Barash et al., 1970;Aziz, 1981a) in the aneuploids. The phenotypic specificity of this aneuploid musculature clearly manifests developmental retardation and instability (Shapiro, 1983) revealing not only the likely course of embryonic myogenesis in chromosomally normal humans (Cihak, 1972, 1977) but also information relevant to ontogenetic and evolutionary changes. The extensor digitorum profundus proprius complex is particularly illustrative of these characteristics of aneuploid musculature. Our examination of the variation of this muscle complex in human aneuploids and between primate genera reveals how normal ontogeny may proceed, as well as the morphological basis for the evolutionary changes in hand structure and function amongst Primates. We also consider the phylogenetic and functional significance of changes in the extensor digitorum profundus proprius with reference to the divergent locomotory and manipulative capabilities and behavior of Primates.     

Comparative myology of the hominoid cranial base. II. The muscles of the prevertebral and upper pharyngeal region

Detailed accounts of the muscular relations and attachments in the prevertebral and upper pharyngeal region of the cranial base of extant hominoids and fossil hominids are described in this paper. Marked differences exist in the cranial attachments of the longus capitis, superior constrictor, tensor palati and levator palati muscles between the three great apes and Homo sapiens. Many of the bony ridges and processes characteristic of the three great apes are also present in the 'gracile' australopithecines. Some of the differences between the muscle markings in this region of the cranial base in certain fossil hominids are discussed in the light of the findings of the dissection study of the three great apes and previous studies of the hominoid cranial base.     

Comparative cranial myology and biomechanics of Plateosaurus and Camarasaurus and evolution of the sauropod feeding apparatus

Sauropodomorpha represents an important group of Mesozoic megaherbivores, and includes the largest terrestrial animals ever known. It was the first dinosaur group to become abundant and widespread, and its members formed a significant component of terrestrial ecosystems from the Late Triassic until the end of the Cretaceous. Both of these factors have been explained by their adoption of herbivory, but understanding the evolution of sauropodomorph feeding has been hampered by the scarcity of biomechanical studies. To address this, the jaw adductor musculature of the basal sauropodomorph Plateosaurus and the sauropod Camarasaurus have been reconstructed. These reconstructions provide boundary conditions for finite element models to assess differences in structural performance between the two taxa. Results demonstrate that Camarasaurus was capable of much greater bite forces than Plateosaurus, due to greater relative adductor muscle mass and shape changes to the mandible. The skull and mandible of Camarasaurus are also ‘stronger’ under static biting. The Plateosaurus mandible appears to compromise structural efficiency and force transmission in order to maintain relatively high jaw closure speed. This supports suggestions of facultative omnivory in basal sauropodomorph taxa. The expanded mandibular symphysis and ‘lateral plates’ of sauropods each lead to greater overall craniomandibular robustness, and may have been especially important in accommodating forces related to asymmetric loading. The functional roles of these characters, and observed general shape changes in increasing skull robustness, are consistent with hypotheses linking bulk‐herbivory with the origin of Sauropoda and the evolution of gigantism.     

The Cell Surface of Trypanosoma musculi Bloodstream Forms. II. Lectin and Immunologic Studies*

SYNOPSIS. Living Trypanosoma musculi bloodstream trypomastigotes were agglutinated specifically with concanavalin A (ConA), wheat germ agglutinin (WGA), soybean agglutinin (SBA), and fucose-binding protein (FBP). The agglutination with these lectins of living cells from which the coat was removed by trypsinization was the same as with intact trypanosomes. Glutaraldehyde or formalin fixation did not affect the results with regard to agglutination with WGA, SBA, and FBP, but lower agglutination with ConA was observed upon fixation. By using a dense iron-dextran marker many fewer ConA marker particles were localized at the fine structural level in the intact than in trypsin-treated trypanosomes. On the basis of the results obtained by agglutination and electron microscopy, it is likely that fixation cross-links intact surface-coat components associated with the ConA binding sites. It is evident from the studies in which lectins were employed that ligands containing α-D-mannose, N-acetylglucosamine, N-acetylgalactosamine, and α-L-fucose are randomly distributed in the outer surface of the pellicular and flagellar membranes of T. musculi trypomastigotes. Results obtained with α-amylase- and dextranase-treated trypanosomes suggested that lectin-binding sugar ligands in the cell surface were not directly associated with α-1,4 or repetitive α-1,6 glucan-bonded polysaccharide moieties. Similar conclusions can be drawn on the basis of neuraminidase treatment with regard to N-acetylated neuraminic acids. After thorough washing, intact, but not trypsin-treated trypomastigotes were agglutinated specifically with antisera against whole mouse serum and against mouse IgG. Evidently, adsorbed constituents of mouse serum are regular components of the T. musculi surface coat. After incubation in dilute whole mouse serum or in mouse IgG solutions, also the trypsinized cells were agglutinated by the 2 antisera. No such results were obtained with trypsinized cells incubated in serum-free buffers. It was concluded that mouse serum proteins were readily readsorbed on, and firmly bound to the trypsinized cells' surfaces. Specific agglutinations were obtained with trypsinized cells after incubation in dilute rat, rabbit, bovine, and human sera and in solutions of rat and rabbit IgG in reactions with the corresponding antisera. It seems, therefore, that the host serum proteins are adsorbed nonspecifically to the cell surface of trypsinized T. musculi bloodstream forms. When examined by electron microscopy, the intact trypomastigotes were covered by an ununiform, slightly granular, fibrillar extracellular coat, applied to the entire outer lamina of the pellicular and flagellar membranes. No indication of such a coat was noted in the trypsinized organisms. Flocculent surface coat-like matrix could, however, be discerned in cells which, after trypsinization, were incubated in various sera.     

Androgens in human evolution. A new explanation of human evolution.

Human evolution consists of chronological changes in gene regulation of a continuous and relatively stable genome, activated by hormones, the production of which is intermittently affected by endogenous and exogenous forces. Periodic variations in the gonadal androgen, testosterone, and the adrenal androgen, dehydroepiandrosterone (DHEA), significantly participated in all hominid transformations. The hominid characteristics of early Australopithecines are primarily a result of increased testosterone. The first significant cold of the early Pleistocene resulted in an increase in DHEA that simultaneously produced Homo and the robust Australopithecines. Subsequent Pleistocene climatic changes and differential reproduction produced changes in DHEA and testosterone ratios that caused extinction of the robust Australopithecines and further changes and continuation of Homo. Changes in testosterone and DHEA produce allometric and behavioral changes that are identifiable and vigorous in modern populations.     

Reflections on the main events and changing factors in the evolution of human behaviour. Present problems of change in human beings from an evolutionary perspective

In this paper the authors analize the main physical and cultural changes of human species from its first moments to the future with an evolutionary perspective. They consider physical items like bipedalism, fire, language or tool production, but also some cultural ones like burials, art or changes on the prehistoric group’s economy, food production, etc.     

Class II genes of the human major histocompatibility complex. Organization and evolutionary relationship of the DR beta genes

The genes of the polymorphic HLA-DR molecules are located within the human major histocompatibility complex. We have studied the HLA-DR genes of an HLA homozygous individual typed to be DR4, Dw4, and DRw53. Fourteen cosmid and phage clones from genomic libraries were isolated and grouped into three clusters comprising a total of 165 kilobases. These clusters contain four DR beta genes. Nucleotide sequence determination showed that two of the genes encode beta chains that carry the DR4 and DRw53 specificities, respectively, while the other two genes are presumably pseudogenes. Comparisons of the nucleotide sequences of all four DR beta genes of the DR4 haplotype show that the genes are extensively similar, approximately 90% in both exons and introns. All four genes are equally similar to each other. These observations are consistent with the notion that the genes arose by duplications that were followed by homogenization through gene conversion. The existence of more than one DR beta gene homologue but only a single DR alpha gene homologue in mouse, rabbit, and cattle suggests that the DR beta gene duplications occurred at or early during mammalian speciation.     

Osteology and myology of the wing of the Emu (Dromaius novaehollandiae), and its bearing on the evolution of vestigial structures

Emus have reduced their wing skeleton to only a single functional digit, but the myological changes associated with this reduction have never been properly described. Moreover, the intraspecific variability associated with these changes has not previously been examined, dissections having been restricted in the past to only one or two individuals. In this paper, the myology and osteology of the Emu wing is described for a sample of five female birds. The Emu showed a marked reduction in the number of muscles in the wing, even compared with other ratites. Many wing muscles showed diversity in structure, origin and insertion sites, number of heads, as well as presence-absence variation. This variability dramatically exceeds that found in flying birds. Evolutionary theory predicts that relaxed selection on vestigial organs should allow more variation to persist in the population, and corresponds to what is observed here. A large amount of fluctuating asymmetry was also detected, indicating reduced canalization of the wing during development.     

Analysis of evolution: evolutionary rates, independence and treeness.

Populations that separate according to a given pattern of fissions (a “tree” of descent) and evolve independently after fissions have a patterned variance—covariance matrix which reflects the history of fissions. Spectral analysis of the matrix can help reconstruct the pattern; multivariate analysis techniques can be used to test if an observed matrix is compatible with a given tree of descent, and with the assumption of constant or of variable evolutionary rates. We call these tests of “treeness” and discuss evolutionary implications, giving also an example on human evolution. The methods can be used more generally to test if a loss of information is involved in using a tree as a formal representation of any particular set of data.