Groundplan Anatomy of the Proboscis of Butterflies (Papilionoidea, Lepidoptera)

The anatomy of the proboscis was studied in representatives of all major subfamilies of Papilionoidea and several outgroup taxa which included Hesperiidae, Hedylidae and Geometroidea. In all species the cross-sectional outline of the tapering proboscis continuously changes from proximal to the tip while the central food canal, formed by the concave medial galeal walls, retains its oval shape. Each galea contains three types of muscles, a branching trachea, nerves, sensilla, and at least one longitudinal septum. We focused on the varying arrangement and distribution of the intrinsic galeal muscles from the basal galeal joint to the tip region. The plesiomorphic condition of the galeal composition of Papilionoidea is regarded to include one basal intrinsic muscle in the basal joint region and two series of intrinsic muscles, i.e. the lateral intrinsic galeal muscles and the median intrinsic galeal muscles, both series extending from the proximal region to the tip region. The plesiomorphic arrangements of the intrinsic muscle series are found in all representatives of Papilionidae, in one species of Lycaenidae (sensu lato), in many Nymphalidae (sensu lato), and in all outgroup species. Three apomorphic character states are distinguished regarding the presence and extension of the median intrinsic galeal muscles. (1) Present up to 35% of the proboscis length and absent distally in Pieridae, Lycaeninae (Lycaenidae), Satyrinae (Nymphalidae), and Danainae (Nymphalidae). (2) Present in the proximal third of the proboscis and again near the tip between 80 and 90% of the proboscis length in the examined Heliconiinae (Nymphalidae). (3) Completely absent, as in one lycaenid species from the subfamily Riodininae.     

Myoanatomy of the marine tardigrade Halobiotus crispae (Eutardigrada: Hypsibiidae)

The muscular architecture of Halobiotus crispae (Eutardigrada: Hypsibiidae) was examined by means of fluorescent‐coupled phalloidin in combination with confocal laser scanning microscopy and computer‐aided three‐dimensional reconstruction, in addition to light microscopy (Nomarski), scanning electron microscopy, and transmission electron microscopy (TEM). The somatic musculature of H. crispae is composed of structurally independent muscle fibers, which can be divided into a dorsal, ventral, dorsoventral, and a lateral musculature. Moreover, a distinct leg musculature is found. The number and arrangement of muscles differ in each leg. Noticeably, the fourth leg contains much fewer muscles when compared with the other legs. Buccopharyngeal musculature (myoepithelial muscles), intestinal musculature, and cloacal musculature comprise the animal's visceral musculature. TEM of stylet and leg musculature revealed ultrastructural similarities between these two muscle groups. Furthermore, microtubules are found in the epidermal cells of both leg and stylet muscle attachments. This would indicate that the stylet and stylet glands are homologues to the claw and claw glands, respectively. When comparing with previously published data on both heterotardigrade and eutardigrade species, it becomes obvious that eutardigrades possess very similar numbers and arrangement of muscles, yet differ in a number of significant details of their myoanatomy. This study establishes a morphological framework for the use of muscular architecture in elucidating tardigrade phylogeny. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Evolutionary developmental studies of cyclostomes and the origin of the vertebrate neck

Because they lack some gnathostome-specific traits, cyclostomes have often been regarded as representing an intermediate state linking non-vertebrate chordates and gnathostomes. To understand the evolutionary origins of the jaw and paired fins, lamprey embryos and larvae have been used as comparative models. The lack of the jaw–neck region is a conspicuous feature specific to cyclostomes; however, the absence of these features has been largely neglected both in evolutionary developmental studies and in the field of classical comparative embryology. This review seeks to develop a possible evolutionary scenario of the vertebrate neck muscles by taking the cucullaris (trapezius) muscle as the focus. By combining the comparative embryology of lampreys and gnathostomes, and considering the molecular-level developmental mechanism of skeletal muscle differentiation, this review argues that the establishment of the vertebrate neck deserves to be called an evolutionary novelty based on the remodeling of mesenchymal components between the cranium and the shoulder girdle, which involves both mesodermal and neural crest cell lineages.     

Heart and craniofacial muscle development: A new developmental theme of distinct myogenic fields

Head muscle development has been studied less intensively than myogenesis in the trunk, although this situation is gradually changing, as embryological and genetic insights accumulate. This review focuses on novel studies of the origins, composition and evolution of distinct craniofacial muscles. Cellular and molecular parallels are drawn between cardiac and branchiomeric muscle developmental programs, both of which utilize multiple lineages with distinct developmental histories, and argue for the tissues' common evolutionary origin. In addition, there is increasing evidence that the specification of skeletal muscles in the head appears to be distinct from that operating in the trunk: considerable variation among the different craniofacial muscle groups is seen, in a manner resembling myogenic specification in lower organisms.     

Comparative dynamics of tropomyosin in vertebrates and invertebrates

Tropomyosin (Tpm) is an extended α-helical coiled-coil homodimer that regulates actinomyosin interactions in muscle. Molecular simulations of four Tpms, two from the vertebrate class Mammalia (rat and pig), and two from the invertebrate class Malacostraca (shrimp and lobster), showed that despite extensive sequence and structural homology across metazoans, dynamic behavior—particularly long-range structural fluctuations—were clearly distinct. Vertebrate Tpms were more flexible and sampled complex, multi-state conformational landscapes. Invertebrate Tpms were more rigid, sampling a highly constrained harmonic landscape. Filtering of trajectories by principle component analysis into essential subspaces showed significant overlap within but not between phyla. In vertebrate Tpms, hinge-regions decoupled long-range interhelical motions and suggested distinct domains. In contrast, crustacean Tpms did not exhibit long-range dynamic correlations—behaving more like a single rigid rod on the nanosecond time scale. These observations suggest there may be divergent mechanisms for Tpm binding to actin filaments, where conformational flexibility in mammalian Tpm allows a preorganized shape complementary to the filament surface, and where rigidity in the crustacean Tpm requires concerted bending and binding.     

More than energy cost: multiple benefits of the long Achilles tendon in human walking and running

Elastic strain energy that is stored and released from long, distal tendons such as the Achilles during locomotion allows for muscle power amplification as well as for reduction of the locomotor energy cost: as distal tendons perform mechanical work during recoil, plantar flexor muscle fibres can work over smaller length ranges, at slower shortening speeds, and at lower activation levels. Scant evidence exists that long distal tendons evolved in humans (or were retained from our more distant Hominoidea ancestors) primarily to allow high muscle–tendon power outputs, and indeed we remain relatively powerless compared to many other species. Instead, the majority of evidence suggests that such tendons evolved to reduce total locomotor energy cost. However, numerous additional, often unrecognised, advantages of long tendons may speculatively be of greater evolutionary advantage, including the reduced limb inertia afforded by shorter and lighter muscles (reducing proximal muscle force requirement), reduced energy dissipation during the foot–ground collisions, capacity to store and reuse the muscle work done to dampen the vibrations triggered by foot–ground collisions, reduced muscle heat production (and thus core temperature), and attenuation of work-induced muscle damage. Cumulatively, these effects should reduce both neuromotor fatigue and sense of locomotor effort, allowing humans to choose to move at faster speeds for longer. As these benefits are greater at faster locomotor speeds, they are consistent with the hypothesis that running gaits used by our ancestors may have exerted substantial evolutionary pressure on Achilles tendon length. The long Achilles tendon may therefore be a singular adaptation that provided numerous physiological, biomechanical, and psychological benefits and thus influenced behaviour across multiple tasks, both including and additional to locomotion. While energy cost may be a variable of interest in locomotor studies, future research should consider the broader range of factors influencing our movement capacity, including our decision to move over given distances at specific speeds, in order to understand more fully the effects of Achilles tendon function as well as changes in this function in response to physical activity, inactivity, disuse and disease, on movement performance.     

同心铲齿象闭合肌组复原及长颌象向短颌象进化过程中头部变化的机制

通过对中新世同心铲齿象(Platybelodon tongxinensis)闭合肌组的复原,讨论了由长颌象向短颌象进化过程中头部形态适应性变化的机制。食性与取食方式变化所引起的下颌变短、下门齿消失,以及由低冠丘脊型、具研磨—剪切功能的臼齿向高冠脊型、具高效水平剪切功能的臼齿的变化是导致头部骨骼和咬合肌组一系列相关适应性变化的主要原因。文章着重讨论、补充了颞肌附着点和延伸方向的变化对于在臼齿齿脊和咬合面积增加的情况下保持臼齿咀嚼面上的压强的功能,并分析了齿脊数增加脊间距减小的功能意义。     

Ascidian larva reveals ancient origin of vertebrate-skeletal-muscle troponin I characteristics in chordate locomotory muscle

Ascidians are protochordates related to vertebrate ancestors. The ascidian larval tail, with its notochord, dorsal nerve cord, and flanking rows of sarcomeric muscle cells, exhibits the basic chordate body plan. Molecular characterization of ascidian larval tail muscle may provide insight into molecular aspects of vertebrate skeletal muscle evolution. We report studies of the Ci-TnI gene of the ascidian Ciona intestinalis, which encodes the muscle contractile regulatory protein troponin I (TnI). Previous studies of a distantly related ascidian, Halocynthia roretzi, showed that different TnI genes were expressed in larval and adult muscles, the larval TnI isoforms having an unusual C-terminal truncation not seen in any vertebrate TnI. Here we show that, in contrast with Halocynthia, Ciona does not have a specialized larval TnI; the same TnI gene that is expressed in the heart and body-wall muscle of the sessile adult is also expressed in embryonic/larval tail muscle cells. Moreover the TnI isoform produced in embryonic/larval muscle is identical to that produced in adult body-wall muscle, i.e., a 182-residue protein with the characteristic chain length and overall structure of vertebrate skeletal muscle TnI isoforms. Phylogenetic analyses indicate that the unique features of Halocynthia larval TnI likely represent derived features, and hence that the vertebrate-skeletal-muscle -like TnI of Ciona is a closer reflection of the ancestral ascidian larval TnI. Our results indicate that characteristics of vertebrate skeletal muscle TnI emerged early in the evolution of chordate locomotory muscle, before the ascidian/vertebrate divergence. These features could be related to a basal chordate locomotory innovation-e.g., swimming by oscillation of an internal notochord skeleton-or they may be of even greater antiquity within the deuterostomes.     

Mapping the Whole-Body Muscle Activity of Hydra vulgaris

1. Download : Download high-res image (206KB)
2. Download : Download full-size image

     

Hominid cultural transmission and the evolution of language

This paper presents the hypothesis that linguistic capacity evolved through the action of natural selection as an instrument which increased the efficiency of the cultural transmission system of early hominids. We suggest that during the early stages of hominization, hominid social learning, based on indirect social learning mechanisms and true imitation, came to constitute cumulative cultural transmission based on true imitation and the approval or disapproval of the learned behaviour of offspring. A key factor for this transformation was the development of a conceptual capacity for categorizing learned behaviour in value terms - positive or negative, good or bad. We believe that some hominids developed this capacity for categorizing behaviour, and such an ability allowed them to approve or disapprove of their offsprings- learned behaviour. With such an ability, hominids were favoured, as they could transmit to their offspring all their behavioural experience about what can and cannot be done. This capacity triggered a cultural transmission system similar to the human one, though pre-linguistic. We suggest that the adaptive advantage provided by this new system of social learning generated a selection pressure in favour of the development of a linguistic capacity allowing children to better understand the new kind of evaluative information received from parents.     

The Importance of Understanding the Nature of Science for Accepting Evolution

Many students reject evolutionary theory, whether or not they adequately understand basic evolutionary concepts. We explore the hypothesis that accepting evolution is related to understanding the nature of science. In particular, students may be more likely to accept evolution if they understand that a scientific theory is provisional but reliable, that scientists employ diverse methods for testing scientific claims, and that relating data to theory can require inference and interpretation. In a study with university undergraduates, we find that accepting evolution is significantly correlated with understanding the nature of science, even when controlling for the effects of general interest in science and past science education. These results highlight the importance of understanding the nature of science for accepting evolution. We conclude with a discussion of key characteristics of science that challenge a simple portrayal of the scientific method and that we believe should be emphasized in classrooms.     

Similarity and diversity in mechanisms of muscle fate induction between ascidian species

Developmental processes can change during evolution at many levels of the ontogeny of an individual. Embryos of solitary ascidians have a largely invariant mode of development, with fixed cleavage patterns and fate maps. Thus the cell lineages and final body plan of the two quite distantly related species considered in this review, Ciona intestinalis and Halocynthia roretzi, are highly similar. However, close comparison of the developmental mechanisms used by these two species provide examples of evolutionary changes and help pinpoint which aspects of development are evolutionarily flexible. Examples of both similarity and diversity are observed in the mechanisms used to generate the full complement of larval muscle. We will describe the changes in muscle-cell lineage, as well as some striking differences in the intercellular signalling pathways used to induce muscle fate. The somewhat surprising conclusion is that in ascidians, as in nematode vulval development, different signalling mechanisms have been adopted to mediate similar interactions between equivalently positioned cells.     

Molluscan muscle systems in development and evolution*

The evolutionary history of the various molluscan muscle systems reflects drastic modifications and reductions as well as true innovations. No less than eight main and independent muscle systems of the Mollusca are described and, based on the current understanding of molluscan phylogeny, their evolutionary histories are outlined.New data on the myogenesis of the Polyplacophora by means of fluorescence‐staining and image analysis by confocal laser scanning microscopy show that the pre‐oral region recapitulates a ‘worm‐grid’, and that the dorso‐ventral musculature passes a stage of multiple seriality as found in adult Solenogastres. Old and new data on bivalves and recent studies on primitive gastropods provide clear evidence that the larval musculature of both groups (and thus possibly of all conchiferans) is entirely independent from the adult condition. The growth of shell‐inserted muscles always necessitates substantial renewal of myocytes which is still poorly understood. Although very promising for phylogenetic purposes, the understanding of the developmental genetics of the various molluscan muscle systems is still in its infancy.     

Science and the Concept of Evolution: From the Big Bang to the Origin and Evolution of Life

The common thread of evolution runs through all science disciplines, and the concept of evolution enables students to better understand the nature of the universe and our origins. “Science and the Concept of Evolution” is one of two interdisciplinary science Core courses taken by Dowling College undergraduates as part of their General Education requirements. The course examines basic principles and methods of science by following the concept of evolution from the big bang to the origin and evolution of life. Case studies of leading scientists illustrate how their ideas developed and contributed to the evolution of our understanding of the world. Evidences for physical, chemical, and biological evolution are explored, and students learn to view the evolution of matter and of ideas as a natural process of change over space and time.     

饥饿和交配对小地老虎飞行肌发育的影响

小地老虎Agrotis ypsilon(Rottemburg)成虫飞行肌的发育常受一些因素影响而发生变化,为探讨饥饿和交配行为对飞行肌发育的影响,通过电子显微镜对雌虫飞行肌(背纵肌)的肌原纤维、线粒体结构进行观察,结果显示:4日龄饥饿雌虫,肌原纤维直径、肌节长度、肌原纤维体积均显著(P<0.05)小于取食的。7日龄饥饿雌虫肌原纤维直径、肌节长度、肌原纤维体积分数较4日龄的差异均不显著(P≥0.05),而7日龄饥饿的肌原纤维直径显著(P<0.05)大于7日龄取食的;羽化10 d后,饥饿雌虫肌节长度显著(P<0.05)大于取食雌虫的,而肌纤维体积分数和线粒体体积分数均却小于后者。7、10、13日龄交配雌虫肌原纤维横切直径分别显著(P<0.05)小于同日龄非交配的;7、10、13日龄交配雌虫肌原纤维体积分数显著(P<0.05)小于非交配的,线粒体体积分数虽然无差异(P≥0.05),但是交配雌虫的早在4日龄便已明显(P<0.05)减小。上述结果表明:正常取食的小地老虎飞行肌4日龄后会发生降解现象;饥饿抑制飞行肌前期发育和中期的降解,而促进成虫末期肌原纤维的分解;交配能促进飞行肌的降解。     

Myo-anatomy of juvenile and adult loxosomatid Entoprocta and the use of muscular body plans for phylogenetic inferences

The anatomy of the muscle bauplan in juvenile buds and adult specimens of the solitary loxosomatid entoprocts Loxosoma nielseni and L. annelidicola was studied by means of fluorescence staining of F-actin and confocal laser scanning microscopy. Although the general myo-anatomy of the body wall shows numerous similarities, both species express significant variations in the arrangement of their pedal muscles. In addition, L. annelidicola alone shows a distinct pair of rectum retractor muscles. Circular muscles are absent in the entire body wall of both species, as well as in previously investigated colonial taxa, which is therefore regarded as basal for Entoprocta. This is in striking contrast to the conditions found in other spiralian or lophotrochozoan taxa. The simple morphology of entoproct tentacle muscles, however, coincides with the phoronid-ectoproct condition and may be due to functional constraints of a simple filter-feeding system. This work shows that variations in the muscular anatomy provide useful characters for systematic analyses on species as well as phylum level and thus allow significant insight regarding metazoan body plan evolution. The phenomenon of phenotypic plasticity and its consequences for phylogenetic interpretations, however, must be carefully considered.     

Rates of evolution: Is there a conflict between neo-darwinian evolutionary theory and the fossil record?

Neo-darwinian and population genetics theory assumes that the necessary and sufficient set of conditions for all genetic, therefore evolutionary, change has been identified. Punctuationalists have assumed the opposite and cite the fossil record as evidence for change too rapid to be explained in neo-darwinian theory. Data is given here to provide estimates of the rate of evolution in hominid fossils, in living populations, and of that rate which would qualify as punctuational in the hominid fossil record. Evolution in living populations is orders of magnitude greater than that found in the fossil record and far greater than necessary to create apparently instantaneous saltations in the fossil record. It is suggested that such saltations may not represent more rapid rates of evolution but, rather, the persistence of evolutionary change in a given direction for a longer than normal period.     

Complex heterochrony underlies the evolution of Caenorhabditis elegans hermaphrodite sex allocation

Hermaphroditic organisms are key models in sex allocation research, yet the developmental processes by which hermaphrodite sex allocation can evolve remain largely unknown. Here we use experimental evolution of hermaphrodite‐male (androdioecious) Caenorhabditis elegans populations to quantify the developmental changes underlying adaptive shifts in hermaphrodite sex allocation. We show that the experimental evolution of increased early‐life self‐fertility occurred through modification of a suite of developmental traits: increased self‐sperm production, accelerated oogenesis and ovulation, and increased embryo retention. The experimental evolution of increased self‐sperm production delayed entry into oogenesis—as expected, given the sequentially coupled production of self‐spermatogenesis and oogenesis. Surprisingly, however, delayed oogenesis onset did not delay reproductive maturity, nor did it trade‐off with gamete or embryo size. Comparing developmental time dynamics of germline and soma indicates that the evolution of increased sperm production did not delay reproductive maturity due to a globally accelerated larval development during the period of self‐spermatogenesis. Overall, heterochrony in gametogenesis and soma can explain adaptive shifts in hermaphrodite sex allocation.     

Troponin T isoforms modulate calcium dependence of the kinetics of the cross-bridge cycle: studies using a transgenic mouse line

Alternative splicing of troponin T (TnT) in striated muscle during development results in expression of different isoforms, with the splicing of a 5(') exon of TnT resulting in the expression of low-molecular-weight basic adult TnT isoforms and high-molecular-weight acidic embryonic TnT isoforms. Although other differences exist, the main differences between cardiac TnT (cTnT) and fast skeletal muscle TnT (fTnT) are in the NH(2) terminus, with fTnT being less acidic than cTnT. A transgenic mouse line expressing chicken fTnT in the heart was used to investigate the functional significance of TnT NH(2)-terminal charge differences on cardiac muscle contractility. The rates of force redevelopment (k(tr)) at four levels of Ca(2+) activation were recorded for skinned left ventricular trabeculae from control and transgenic mice. The k(tr) vs Ca(2+) relationship was different in control mice and transgenic mice, suggesting that the structure of TnT, and possibly the NH(2)-terminal region, is involved in determining the kinetics of cross-bridge cycle. These results suggest that isoform shifts in TnT may be an important molecular mechanism for determining the Ca(2+) dependence of cardiac muscle contractility.     

Higher rates of sex evolve under K‐selection

The geographical distribution of sexual and related asexual species has been suggested to correlate with habitat stability; sexual species tend to be in stable habitats (K‐selection), whereas related asexual taxa tend to be in unstable habitats (r‐selection). We test whether this broad‐scale pattern can be re‐created at a microevolutionary scale by experimentally evolving populations of facultatively sexual rotifers under different ecological conditions. Consistent with the pattern in nature, we find that the rate of sex evolves to lower levels in the r‐selected than in K‐selection environments. We consider several different explanations for these results.