Particle retention and flow in the pharynx of the enteropneust worm Harrimania planktophilus: the filter-feeding pharynx may have evolved before the chordates

An investigation of the feeding behavior of the acorn worm Harrimania planktophilus suggests a novel form of enteropneust feeding with significant phylogenetic implications. H. planktophilus is a holoinfaunal worm that feeds on deposited sediments, and filter feeds on suspended particles in interstitial pore water. To visualize the particle retention behavior involved in filter feeding, adult animals were held in chilled seawater under low light and fed food coloring and fluorescent particles. The behavior was recorded by videography. Most particles ingested were drawn into the mouth by an incurrent flow created by cilia on the pharyngeal bars and without the aid of mucus. Particles that passed freely through the gill pores averaged 3.04 microm, whereas particles retained in the gut and defecated in the feces averaged 13.9 microm. Food coloring entered the mouth and was pumped through the pharynx at a rate of 0.5-2.0 mm/s. There is no evidence of an endostyle or mucus-net capture mechanism in H. planktophilus, but instead particles are filtered and manipulated by a dense covering of cilia on the pharyngeal gill bars. This study suggests that the filter-feeding pharynx is not an innovation of the chordates, but evolved prior to the evolutionary divergence of the hemichordate-echinoderm clade from the chordates.     

Pelagic eggs and larvae of Coelorinchus kishinouyei (Gadiformes: Macrouridae) collected from Suruga Bay, Japan

Pelagic eggs and larvae of the macrourid fish Coelorinchus kishinouyei, collected from Suruga Bay, southern Japan and subsequently identified by 16S rRNA gene nucleotide sequences, are described. The spherical eggs, 1.18–1.31 mm in diameter, contained a single oil globule, 0.28–0.33 mm in diameter, and had hexagonally patterned ornamentation on the chorion, 0.017–0.022 mm in width. Melanophores were present on the embryo, yolk and oil globule after the blastopore had closed. Within 1 day after hatching, the body axis of the yolk-sac larvae was bent slightly at the anterior trunk region. During this stage many melanophores formed on the head, trunk, tail, yolk and oil globule, along with small irregular wrinkles on the dorsal and ventral finfolds. Pelagic eggs (after the caudal end of the embryo had detached from the yolk) and yolk-sac larvae also developed xanthophores on the embryo and yolk, and head, trunk, dorsal and ventral finfolds just before tail tip, and yolk, respectively. The pelagic larvae had a short tail, stalked pectoral-fin base and no elongate first dorsal and pelvic-fin rays. Three clusters of melanophores were present on the tail (anterior two embedded to muscle and one just before tail tip subsequently lost with development) and a cluster around the anus (beyond 3.9 mm head length). Nucleotide sequence analyses of comparative adult specimens appeared to confirm a previous proposal that C. productus is a junior synonym of C. anatirostris.     

Early stages of spinal ganglion formation during tail regeneration in the newt, Notophthalmus viridescens

Stages in the development of sensory ganglia in the regenerating newt tail after amputation are described by taking advantage of the rostrocaudal developmental gradient of the regenerating tail. A series of ganglia, beginning at the tip of the regenerate and progressing rostrally, were examined. Eight-week regenerates were used because they showed the most complete array of stages. The first recognizable ganglia appear as small clusters of cells sitting dorsally on the already established ventral roots. The cluster of ganglionic cells steadily expands with the addition of many new cells. Signs of cell differentiation within the ganglion precede the formation of the dorsal root rudiment, which assumes several different configurations but most commonly enters the cord close to the ventral root. Our material suggests that ganglion precursor cells originate in the ventral region of the developing spinal cord and migrate out of the cord by travelling along the ventral root until, at a suitable distance from the cord, they halt, proliferate, and eventually differentiate. In the regenerate, we saw no evidence of neural crest cells--such as those that give rise to ganglia in the trunk region during development--forming at the dorsal region of the regenerated neural tube. Nor was there any morphological evidence of mesenchymal contribution to the ganglion cell clusters.     

Early ontogeny of the commercially valuable fish red-bellied pacu Piaractus brachypomus (Characiformes,Serrasalmidae) from the Amazon,Brazil

The initial development of the red-bellied pacu Piaractus brachypomus is described using morphological, meristic and morphometric characteristics. A total of 127 individuals were analysed (47 in the yolk-sac, 35 in pre-flexion, five in flexion, 20 in post-flexion and 20 in juvenile) with standard length varying between 2.92 and 48.61 mm. The larvae are born poorly developed and have a discoidal yolk at ~6.33 mm standard length. During early ontogeny, the mouth passes from terminal to subterminal and the anal opening reaches the vertical line over the midline region of the body. There are changes in body shape from long and moderate to deep, head length from small to large, and eye diameter from moderate to large. Dendritic chromatophores were present in the ventral, dorsal and upper part of the swim bladder in the early larval stages. Rounded spots are evident all over the body in juveniles. The total number of myomeres ranges from 39 to 41 (20–23 pre-anal, 17–20 post-anal). Through the morphometric relationships, it was evidenced that the greatest changes during the initial ontogeny of P. brachypomus occur in the transition from the post-flexion stage to the juvenile period, indicating changes in behaviour, foraging and physiology.     

Neosteinernema longicurvicauda n. gen., n. sp. (Rhabditida: Steinernematidae), a Parasite of the Termite Reticuldermes flavipes (Koller)

A nematode isolated from the termite Reticulitermes flavipes (Koller) was identified and described as a new genus and species, Neosteinernema longicurvicauda. Primary distinguishing characters, by contrast to members of the genus Steinernema, were females having prominent phasmids, a curved tail longer than the body width at the anus, a spiral shape in juvenile-bearing females, and juveniles becoming infective-stage juveniles before emerging from the female; males having prominent phasmids, a digitate tail tip, a characteristic shape of the spicules (foot-shaped with a hump on the dorsal side), and 13-14 pairs of genital papillae, with eight pairs preanal; and infective juveniles having prominent phasmids and a filiform curved tail as long as the esophagus. Adult nematodes are found outside the termite cadaver. Diagnosis of the family Steinernematidae was emended to accommodate the new species.     

The postanal tail of the enteropneust Saccoglossus kowalevskii is a ciliary creeping organ without distinct similarities to the chordate tail

Stach, T. and Kaul, S. 2011. The postanal tail of the enteropneust Saccoglossus kowalevskii is a ciliary creeping organ without distinct similarities to the chordate tail. —Acta Zoologica (Stockholm) 92 : 150–160. The postanal tail of chordates is one of the key characters in chordate evolution and it has been suggested to be homologous to the postanal tail of harrimaniid enteropneusts. We present electron microscopic data of the ontogeny of the postanal tail in the enteropneust Saccoglossus kowalevskii. The postanal tail develops as a ventral posterior allometric outgrowth with a ventral extension of the telotroch. Transmission electron microscopy of serial sections reveals the epidermal organization of the postanal tail with the exception of short, bilaterally symmetric extensions of the paired metacoels. The epidermis cells are connected by apical junctions, rest basally on the extracellular matrix surrounding the mesoderm, and possess a basiepidermal nerve net. The ventral cells in the postanal tail are multiciliated and used for creeping. Dorsal cells are monociliated with numerous microvilli. Two types of glandular cells are present among the epidermis cells. The mesoderm cells contain myofilaments. We were unable to detect anatomical structures similar to the ones present in the postanal locomotory tail of chordates, such as notochord, neural tube, or endodermal strand. Thus, results of our anatomical study do not support homology of the postanal chordate tail and the postanal tail of harrimaniid enteropneusts.     

Dorsoventral axis inversion: Enteropneust anatomy links invertebrates to chordates turned upside down

The relationships between chordates with their dorsal nerve cord and other animal groups remain unclear. The hemichordata, specifically the enteropneusta (acorn worms), have been considered a sister group to the chordata. Enteropneusts combine various chordate features (e.g. lateral gill openings, dorsal nerve cord) with features that are usually associated with gastroneuralian invertebrates (e.g. dorsal heart, circumenteric nerve ring, ventral nerve cord). Here we analyse various morphological and functional characteristics that enteropneusts share with either invertebrates or chordates in the light of our recent proposal that the chordata may derive – by bodily dorsoventral inversion – from a gastroneuralian ancestor. We show that many seemingly non-chordate features of enteropneusts will align with similar features in the chordates – provided that we compare the ventral side of an enteropneust to the dorsal side of a chordate. This inversion proposes several interesting and new putative homologies between enteropneusts and acranian chordates, such as between their epibranchial ridge/endostyle (later thyroid gland), their postanal tails, atrial walls, and also between the chordates' dorsal notochord and the enteropneusts' posteroventral pygochord. Significantly, positional homology between notochord and pygochord is also supported by the expression domains of Brachyury orthologs in vertebrates and invertebrates: a Brachyury ortholog is active in the postero ventral mesoderm in Drosophila and in the dorsal mesoderm in chordates. In conclusion, we propose that the anatomy of enteropneusts may serve as a conceptual 'missing link' between gastroneuralian invertebrates and notoneuralian chordates. We discuss whether the enteropneust's dorsoanterior nervous centre plus their ventral trunk cord then corresponds to brain and dorsal nerve cord in the chordata.     

弓形革囊星虫组织结构和超微结构观察

弓形革囊星虫(Phascolosoma arcuatum)是近年兴起的水产养殖品种,目前已形成一定养殖规模,但是其生物学特性相关的研究只有零星报道。本文主要采用石蜡切片苏木精-伊红(H.E)染色和扫描电镜方法对弓形革囊星虫的整体形态和各主要器官的结构进行研究。结果表明,弓形革囊星虫主要由吻部和躯干部构成,躯干部有乳突分布;吻部由触手和项器构成,伸长可达躯干部的两倍,具有感光性。其消化系统由口、食道、肠、肛门四个部分构成,不同部分的肌肉组织存在明显差异;收吻肌一端与吻相连,另一端分成4支,分别连于背部两侧(背收吻肌)和腹部两侧(腹收吻肌);食道贴于收吻肌,后连接肠。肠分为肠下回环与肠上回环,肠上回环与肠下回环缠绕盘旋,肠上回环后接直肠。直肠具有盲囊,后连接肛门;两条肾管分布在腹侧。     

Description of the Kona Coffee Root-knot Nematode,Meloidogyne konaensis n. sp.

Meloidogyne konaensis n. sp. is described from coffee from Kona on the island of Hawaii. The perineal pattern of the female is variable in morphology, the medial lips of the female are divided into distinct lip pairs, and the excretory pore is 2-3 stylet lengths from the base of the stylet. Mean stylet length is 16.0 μm, and the knobs gradually merge with the shaft. The knobs are indented anteriorly and rounded posteriorly and the dorsal esophageal gland orifice (DEGO) is long, 3.5-7 μm. The morphology of the stylet of the male is the most useful diagnostic character, with 6-12 large projections protruding from the shaft. One medial lip may be divided into distinct lip pairs. A large intestinal caecum often extends nearly to the level of the DEGO. Mean juvenile length is 502 μm, mean stylet length is 13.4 μm, and mean tail length is 58 μm. The tail may be distinctly curved ventrally and the phasmids are located in the ventral incisure about one anal body width posterior to the anus.     

Saccoglossus testa from the Mazon Creek fauna (Pennsylvanian of Illinois) and the evolution of acorn worms (Enteropneusta: Hemichordata)

The limited fossil record of enteropneust hemichordates (acorn worms) and the few external features that distinguish the four families have provided a challenge to our understanding of the evolution of the group and their various feeding adaptations. The middle Pennsylvanian Saccoglossus testa sp. nov. from the Mazon Creek, Westfalian D Carbonate Formation, Francis Creek Shale of northern Illinois provides evidence for the exploitation of surface sediments. Saccoglossus testa has a long proboscis characteristic of the extant genus Saccoglossus, a specialist in surface deposit feeding. The collar is as long as it is wide. The anterior trunk lacks a distinctively wide branchial region. These three features distinguish it from its sympatric enteropneust species Mazoglossus ramsdelli Bardack that has a proboscis characteristic of an infaunal deposit feeder. It is the seventh known species of fossil enteropneust, including a resting trace of a Lower Triassic fossil that has collar lips that characterize the extant deep‐sea family Torquaratoridae, and which represents a second parallel evolution of surface deposit feeding. An analysis of the seven fossils shows that the earliest Enteropneusta had a relatively simple harrimaniid‐like body plan, and that the spengelid, the torquaratorid and lastly the most complex ptychoderid body plan appeared in that chronological order.     

16细胞期爪蟾胚胎背方与腹方裂球的发育能力

在两栖类爪蟾胚胎发育中,由受精引起的皮层转动造成了受精卵的背腹极性。为了研究受精卵细胞质的不均一分布对胚胎体轴形成的影响,我们进行了16细胞期动物极背、腹方裂球的外植和异位移植实验。16细胞期的动物极背方裂球在外植和移植到腹方位置后都表现出背方特征,如外植块培养到原肠中期时伸长,背方裂球在移植到腹方后引发第二体轴等;而16细胞期动物极腹方裂球移植到背方后其发育命运则遵循背方裂球的命运,参与背方结构的形成。我们认为在16细胞期,动物极背、腹方的裂球由于包含着不同的卵质,因而在发育能力上已经具有背、腹的差异。     

Segmental organisation of the tail region in the embryo of Drosophila melanogaster

Summary The segmental organisation of the tail region in the embryo of Drosophila melanogaster, which is defined here as the epidermal region posterior to the boundary between abdominal segments A7 and A8, has been investigated by means of ultraviolet (UV) laser fate-mapping and phenotypic analysis of embryonic mutants that alter the segmental pattern of the larval cuticle. Wild-type embryos were irradiated in the presumptive tail region with a UV- laser microbeam of 20 m diameter at the blastoderm stage. The ensuing defects were scored in the cuticle pattern of the tail region of the first-instar larva, which is described in detail in this paper. The spatial distribution of defect frequencies was used to construct a blastoderm fate-map of the cuticle structures of the larval tail region. The segmental origin of the larval tail structures was inferred from the phenotypic analysis of segmentation and homoeotic mutants, which revealed pattern repetition throughout the embryonic tail region corresponding to four segment anlagen, A8 to A11, and a non-segmental telson. These data enabled the transformation of the blastoderm fate-map of cuticle structures into a map of tail segment anlagen. The tail anlage occupies about 10% of the egg length (EL), bounded by segment A7 anteriorly at 20% EL and by the proctodaeum posteriorly at 10% EL, as measured from the posterior pole. The anlagen of segments A8 and A9 appear to be narrow dorso-ventral strips of blastoderm cells similar to the anlagen of the trunk segments, whereas the anlagen of A10 and A11 are smaller and produce fewer pattern elements. The telson is represented in the cuticle by the tuft which derives from a very dorsal posterior position. The antero-posterior axis of the entire tail anlage appears curved upward posteriorly. Differences in the mode of development between tail and trunk segments are discussed, as are similarities of larval and imaginal tail development in Drosophila. Comparison with tail development in other insects suggests that, during evolution, the transition from semi-long-germ to long-germ development modified the organisation of the tail region without affecting its primary subdivision into metameric units.     

The development of social behaviour in the vole (Microtus agrestis)

Juvenile voles of the species Microtus agrestis investigated the bodies of other juveniles with the nose, by touching the dorsal surface from snout to tail. Play movements often followed such nosing contacts and augmented further contacts. The most important scent stimulating play was produced by the skin at the back of the head. The secretion of the nose and mouth region may convey sexual identity, the rump region and feet individual identity.
Young born in the spring and early summer often interacted and played, the autumn young seldom. The behaviour of overwintering sexually immature adults was similar to that of the autumn juveniles. It is suggested that this seasonal difference in the behaviour of juveniles has consequences for subsequent mature males, and that this developmental factor may have an important effect on the annual population cycle.     

A new species of lentic breeding Korean salamander of the genus Hynobius (Amphibia, Urodela)

A new species of salamander, Hynobius yangi, is described from Kori (=Hyoam-ri), Busan-shi, southeastern Korea. It is a lentic breeder belonging to the H. nebulosus species group, and has long been confused with another Korean species H. leechii. The new species, however, is genetically substantially differentiated from Korean and Japanese relatives. Hynobius yangi is morphologically very similar to H. leechii, but could be differentiated from it by the tail shape, degree of limb separation, shape of vomerine teeth series, shape of egg sac, and dorsal coloration.     

Organization of the superficial neuromast system in goldfish, Carassius auratus

Distribution, morphology, and orientation of superficial neuromasts and polarization of the hair cells within superficial neuromasts of the goldfish (Carassius auratus) were examined using fluorescence labeling and scanning electron microscopy. On each body side, goldfish have 1,800-2,000 superficial neuromasts distributed across the head, trunk and tail fin. Each superficial neuromast had about 14-32 hair cells that were arranged in the sensory epithelium with the axis of best sensitivity aligned perpendicular to the long axis of the neuromast. Hair cell polarization was rostro-caudal in most superficial neuromasts on trunk scales (with the exception of those on the lateral line scales), or on the tail fin. On lateral line scales, the most frequent hair cell polarization was dorso-ventral in 45% and rostro-caudal in 20% of the superficial neuromasts. On individual trunk scales, superficial neuromasts were organized in rows which in most scales showed similar orientations with angle deviations smaller than 45 degrees . In about 16% of all trunk scales, groups of superficial neuromasts in the dorsal and ventral half of the scale were oriented orthogonal to each other. On the head, most superficial neuromasts were arranged in rows or groups of similar orientation with angle deviations smaller than 45 degrees . Neighboring groups of superficial neuromasts could differ with respect to their orientation. The most frequent hair cell polarization was dorso-ventral in front of the eyes and on the ventral mandible and rostro-caudal below the eye and on the operculum.     

Test cell migration and tunic formation during posthatching development of the larva of the ascidian, Ciona intestinalis

Morphological changes in the tunic layers and migration of the test cells during swimming period in the larva of the ascidian, Ciona intestinalis , were observed by light and electron microscopy. The swimming period was divided into three stages. In stage 1, further formation of juvenile tunic layer started only in the larval trunk and neck region. In stage 2, the layer became swollen in the ventral and dorsal sides of the neck region and in stage 3, the swelling expanded backward. Concomitantly with these changes, the outermost larval tunic layer (outer cuticular layer), which had been formed before hatching, also swelled in the neck region in stage 2 and formed two humps in stage 3, although the layer did not change in the tail region during the swimming period. Test cells that were present over the entire larval tunic layer in stage 1 began to move from the surface of the fin toward that of the side of the body in stage 2, and finally gathered to form six bands running radially from the anterior end to the posterior end of the trunk region and aligned along the lateral sides of body in the tail region in stage 3. In electron microscopic observations, pseudopodia protruding from the test cells invaded the larval tunic, following which they extended proximate to the juvenile tunic in the trunk region. In the tail region, which had no juvenile tunic layer as that described, the pseudopodia invaded and remained adjacent to the surface of the epidermis or the sensory cilia protruded from the epidermis. Metamorphosis of the larvae, further tunic formation, degradation of adhesive papilla, attachment of larva to the substratum and tail resorption commenced after these morphological changes occurred. The possible role of the test cells in metamorphosis is discussed.     

The Fine Structure of Some Blood Vessels of the Earthworm, Eisenia foetida

The fine structure of the main dorsal and ventral circulatory trunks and of the subneural vessels and capillaries of the ventral nerve cord of the earthworm, Eisenia foetida, has been studied with the electron microscope. All of these vessels are lined internally by a continuous extracellular basement membrane varying in thickness (0.03 to 1 µ) with the vessel involved. The dorsal, ventral, and subneural vessels display inside this membrane scattered flattened macrophagic or leucocytic cells called amebocytes. These lie against the inner lining of the basement membrane, covering only a small fraction of its surface. They have long, attenuated branching cell processes. All of these vessels are lined with a continuous layer of unfenestrated endothelial cells displaying myofilaments and hence qualifying for the designation of "myoendothelial cells." The degree of muscular specialization varies over a spectrum, however, ranging from a delicate endowment of thin myofilaments in the capillary myoendothelial cells to highly specialized myoendothelial cells in the main pulsating dorsal blood trunk, which serves as the worm's "heart" or propulsive "aorta." The myoendothelial cells most specialized for contraction display well organized sarcoplasmic reticulum and myofibrils with thick and thin myofilaments resembling those of the earthworm body wall musculature. In the ventral circulatory trunk, circular and longitudinal myofilaments are found in each myoendothelial cell. In the dorsal trunk, the lining myoendothelial cells contain longitudinal myofilaments. Outside these cells are circular muscle cells. The lateral parts of the dorsal vessels have an additional outer longitudinal muscle layer. The blood plasma inside all of the vessels shows scattered particles representing the circulating earthworm blood pigment, erythrocruorin.     

Ultrastructure of hatchling chaetognaths (Ferosagitta hispida): Epithelial arrangement of the mesoderm and its phylogenetic implications

The trunk and tail mesoderm of hatchling chaetognaths consists of a simple myoepithelium containing four stereotypically arranged cell types, each matching in position a specific adult tissue. The trunk mesoderm includes lateral cells, longitudinal muscle cells, dorsal and ventral medial cells, and peri-intestinal cells. These correspond, respectively, to the lateral fields, longitudinal body wall muscles, dorsal and ventral perimysial cells, and periintestinal muscles of adults. Because the developing intestine does not extend into the tail, tail cells equivalent in position to peri-intestinal cells in the trunk are designated mesenterial cells. Numerous small spaces situated among the apices of hatchling mesodermal cells have the same position relative to surrounding cells as both the coelomic cavities of early embryos and the adult body cavities. We infer that these spaces in hatchlings expand and coalesce to form the definitive adult body cavities, and that these spaces and the adult body cavities derive from the embryonic coeloms. Because hatchlings lack mesenchymal mesoderm, we infer that all adult mesodermal tissues develop by elaboration of the coelomic lining of hatchlings. Because hatchlings lack cells corresponding to the squamous peritoneocytes overlying the body wall muscles of adults, we conclude that peritoneocytes are specialized adult cells that are not equivalent to cells of the embryonic coelomic lining. Finally, hatchlings contain a complete trunk/tail septum. This observation contradicts reports that this septum forms several days after hatching. It also weakens arguments that chaetognaths are bimeric rather than trimeric. © 1994 Wiley-Liss, Inc.     

Ontogeny of the collar cord: Neurulation in the hemichordate Saccoglossus kowalevskii

The chordate body plan is characterized by a central notochord, a pharynx perforated by gill pores, and a dorsal central nervous system. Despite progress in recent years, the evolutionary origin of each of theses characters remains controversial. In the case of the nervous system, two contradictory hypotheses exist. In the first, the chordate nervous system is derived directly from a diffuse nerve net; whereas, the second proposes that a centralized nervous system is found in hemichordates and, therefore, predates chordate evolution. Here, we document the ontogeny of the collar cord of the enteropneust Saccoglossus kowalevskii using transmission electron microscopy and 3D‐reconstruction based on completely serially sectioned stages. We demonstrate that the collar cord develops from a middorsal neural plate that is closed in a posterior to anterior direction. Transversely oriented ependymal cells possessing myofilaments mediate this morphogenetic process and surround the remnants of the neural canal in juveniles. A mid‐dorsal glandular complex is present in the collar. The collar cord in juveniles is clearly separated into a dorsal saddle‐like region of somata and a ventral neuropil. We characterize two cell types in the somata region, giant neurons and ependymal cells. Giant neurons connect via a peculiar cell junction that seems to function in intercellular communication. Synaptic junctions containing different vesicle types are present in the neuropil. These findings support the hypotheses that the collar cord constitutes a centralized element of the nervous system and that the morphogenetic process in the ontogeny of the collar cord is homologous to neurulation in chordates. Moreover, we suggest that these similarities are indicative of a close phylogenetic relationship between enteropneusts and chordates. J. Morphol., 2010. ©2010 Wiley‐Liss, Inc.     

Location of the Phasmids on Infective Juveniles of Steinernema glaseri

Scanning electron microscopy revealed the location of the phasmids on infective juveniles of Steinernema glaseri. The phasmids are located about 40% of the tail length posterior to the anus and are at or near the same level. Instead of being in the center of the lateral fields, they are located just ventral to the lateral fields, or interrupting the ventral-most lateral ridge. The phasmids were covered often by an exudate.