What are the roles of retinoids,other morphogens,and Hox genes in setting up the vertebrate body axis?

This article is concerned with the roles of retinoids and other known anterior–posterior morphogens in setting up the embryonic vertebrate anterior–posterior axis. The discussion is restricted to the very earliest events in setting up the anterior–posterior axis (from blastula to tailbud stages in Xenopus embryos). In these earliest developmental stages, morphogen concentration gradients are not relevant for setting up this axis. It emerges that at these stages, the core patterning mechanism is timing: BMP‐anti BMP mediated time space translation that regulates Hox temporal and spatial collinearities and Hox‐Hox auto‐ and cross‐ regulation. The known anterior–posterior morphogens and signaling pathways––retinoids, FGF's, Cdx, Wnts, Gdf11 and others––interact with this core mechanism at and after space–time defined “decision points,” leading to the separation of distinct axial domains. There are also other roles for signaling pathways. Besides the Hox regulated hindbrain/trunk part of the axis, there is a rostral part (including the anterior part of the head and the extreme anterior domain [EAD]) that appears to be regulated by additional mechanisms. Key aspects of anterior–posterior axial patterning, including: the nature of different phases in early patterning and in the whole process; the specificities of Hox action and of intercellular signaling; and the mechanisms of Hox temporal and spatial collinearities, are discussed in relation to the facts and hypotheses proposed above.     

Gene expression suggests conserved mechanisms patterning the heads of insects and myriapods

Segmentation, i.e. the subdivision of the body into serially homologous units, is one of the hallmarks of the arthropods. Arthropod segmentation is best understood in the fly Drosophila melanogaster. But different from the situation in most arthropods in this species all segments are formed from the early blastoderm (so called long-germ developmental mode). In most other arthropods only the anterior segments are formed in a similar way (so called short-germ developmental mode). Posterior segments are added one at a time or in pairs of two from a posterior segment addition zone. The segmentation mechanisms are not universally conserved among arthropods and only little is known about the genetic patterning of the anterior segments. Here we present the expression patterns of the insect head patterning gene orthologs hunchback (hb), orthodenticle (otd), buttonhead-like (btdl), collier (col), cap-n-collar (cnc) and crocodile (croc), and the trunk gap gene Krüppel (Kr) in the myriapod Glomeris marginata. Conserved expression of these genes in insects and a myriapod suggests that the anterior segmentation system may be conserved in at least these two classes of arthropods. This finding implies that the anterior patterning mechanism already existed in the last common ancestor of insects and myriapods.     

Description of two new miniature species of Hylopanchax Poll & Lambert, 1965 (Cyprinodontiformes: Procatopodidae), from northeastern Gabon,with an updated diagnosis of the genus based on morphology,colouration and osteology

Two new species of the lampeye genus Hylopanchax are described from the Ivindo River basin in the Ogowe River drainage. Hylopanchax multisquamatus, new species, and Hylopanchax thysi, new species, differ from congeners by the presence of a hyaline urogenital male papilla with small black spots and a dark-brown reticulate pattern on the flanks of both males and females in preserved specimens. Hylopanchax multisquamatus is distinguished from congeners by the number of scales on the mid-longitudinal series (27–30 vs. 19–26, respectively) and by the relative anterior/posterior flank scale depth ratio (140%–150% vs. 170%–220%). Hylopanchax thysi is distinguished from all other congeners, except Hylopanchax paucisquamatus, by the presence of vertebrae (30 vs. 31–33) and is further distinguished from H. multisquamatus by the presence of a deeper caudal peduncle and much larger anterior flank scales. It is distinguished from H. paucisquamatus by the presence of a hyaline urogenital male papilla with small black spots and a dark-brown reticulate pattern on the flanks of both males and females in preserved specimens. Osteological data of Hylopanchax are presented for the first time, and an updated diagnosis based on external morphology, colouration pattern and osteology is provided. An osteological comparison with closely related species belonging to the genera Procatopus, Hypsopanchax and “Hypsopanchax” is presented. (a) A truncate and slightly downward-directed anterior process of the angulo-articular and (b) a guitar-shaped lachrymal with both its anterior and posterior margins sharply curved are here considered as diagnostic features of Hylopanchax.     

Molecular studies of hemichordate development: a key to understanding the evolution of bilateral animals and chordates

SUMMARY Using the Hawaiian acorn worm, Ptychodera flava, we began molecular studies on the development of hemichordates, a phylum previously unstudied at this level. Here we review results garnered from the examination of a few specific genes selected to help understand the evolution of vertebrate structures. These studies suggest new ideas about the evolution of developmental mechanisms in the deuterostomes. In a seminal observation, we noted an unexpected zone of expression of the Brachyury gene in the early anterior embryonic ectoderm where the mouth will form. Typically, the Brachyury gene is closely linked to development of the notochord and is expressed around the blastopore and in the posterior mesoderm in most animals. This first expression of Brachyury at the blastopore may represent a regulatory program associated with organizing the original animal head and gut opening, as suggested by the expression of Brachyury during hypostome formation in hydra. We believe that the anterior expression of Brachyury in deuterostomes represents the cooption of the program for organizing the original animal gut opening to form the deuterostome mouth. Recent data from the trochophore larva of a polychaete show that an anterior zone of expression of Brachyury is produced in this protostome by splitting of the Brachyury field during the formation of a gut with a mouth and anus by the lateral fusion of the sides of the blastopore. The ability to initiate independently a secondary regulatory program to organize the new mouth leading to an anterior field of Brachyury expression may be a signal event in the evolution of the deuterostomes. We also noted that the P. flava homolog of T‐brain/Eomes, a gene closely related by sequence and expression around the blastopore to Brachyury and associated with development of the vertebrate brain, also exhibits early posterior expression around the blastopore and a field of de novo anterior ectoderm expression during later embryogenesis. The tissue in the zone of de novo anterior ectoderm expression of Pf‐Tbrain produces the apical organ, a larval neural structure that has been touted as an evolutionary precursor of the chordate dorsal brain. The gene regulatory mechanisms responsible for initiating the anterior zone of de novo expression of T‐brain may represent a cooption to specify early neuroectoderm of the regulatory program evolved first to drive anterior Brachyury expression for deuterostome mouth formation. It will be interesting to examine the possibilities that an ability to initiate the de novo anterior expression of the program that includes T‐brain may be a key event in the evolution of the developmental mechanisms leading to the chordate dorsal nervous system.     

Choanoflagellates (Sarcomastigophora, Protozoa) from the Coastal Waters of Taiwan and Japan. I. Three New Species

Three new acanthoecidaean species collected from the coastal waters of Taiwan and Japan are described: Acanthocorbis camarensis n. sp. resembles Acanthocorbis unguiculata in lorica morphology, but differs in having regularly arranged longitudinal and transverse costae at the anterior lorica chamber, and in lacking a nail at the apical end of anterior spine; Diaphanoeca spiralifurca n. sp. is characterized by the spiral arrangement of the costal strips in the posterior half of the lorica chamber and is closely related to Diaphanoeca grandis; Stephanoeca supracostata n. sp. is closely related to Stephanoeca elegans, but differs in having an additional transverse costa at the anterior lorica chamber.     

Effects of combined centrifugation and ultraviolet irradiation of eggs on pattern formation in Chironomus embryos

Summary

Combined mild centrifugation and uv irradiation of Chironomus embryos modified the developmental types expected from centrifugation alone, somewhat differently from the combined strong centrifugation and uv irradiation of Smittia embryos. The modifications changed with the stages irradiated. The change caused by anterior irradiation may depend on whether or not a part of the cytoplasmic zone is irradiated simultaneously with the anterior yolky end; because most of the cytoplasm lies in the posterior half of egg at early irradiation, while the tip of the cytoplasm redistributes near the anterior end by the late irradiation. Early uv irradiation of the anterior end of centrifuged eggs, causing the formation of a double abdomen (DA) or an inverted embryo, is not photoreversible, while the uv damage to the anterior end of uncentrifuged eggs, inducing DA, is. These facts suggest that there is another photoirreversible uv target in addition to the photoreversible target for DA induction or the anterior determinant shown in Smittia. Other changes, such as the induction of a double cephalon by late irradiation of the centrifuged egg, are photoreversible, but in an unusual way in that the level of photorecovery is similar to the result of incubation in the dark after early irradiation, and not to that of the centrifuged controls. These modified results were then compared with those for Smittia embryos.     

Steganoderma szidati n. sp. (Trematoda: Zoogonidae) from Galaxias maculatus (Jenyns) and G. platei Steindachner in Patagonia,Argentina

Steganoderma szidati n. sp. is described from the intestine of two freshwater fishes, Galaxias maculatus and G. platei (Galaxiidae), from Andean lakes in Patagonia, Argentina. This freshwater zoogonid species fits in the genus Steganoderma (sensu Bray, 1987) because of the length of the caeca and the position of the vitelline follicles. The new species is characterised by possesing 6-13 vitelline follicles situated between the anterior margin of the ventral sucker and the poterior margin of the testis. The gonads are in the anterior hindbody and the ovary is anterior to the right testis. The cirrus has two conspicuous spines at its distal end, and the seminal vesicle always exhibits a constriction. The excretory vesicle never reaches the level of the posterior margin of the testes.     

Bmpr1a signaling plays critical roles in palatal shelf growth and palatal bone formation

Cleft palate, including submucous cleft palate, is among the most common birth defects in humans. While overt cleft palate results from defects in growth or fusion of the developing palatal shelves, submucous cleft palate is characterized by defects in palatal bones. In this report, we show that the Bmpr1a gene, encoding a type I receptor for bone morphogenetic proteins (Bmp), is preferentially expressed in the primary palate and anterior secondary palate during palatal outgrowth. Following palatal fusion, Bmpr1a mRNA expression was upregulated in the condensed mesenchyme progenitors of palatal bone. Tissue-specific inactivation of Bmpr1a in the developing palatal mesenchyme in mice caused reduced cell proliferation in the primary and anterior secondary palate, resulting in partial cleft of the anterior palate at birth. Expression of Msx1 and Fgf10 was downregulated in the anterior palate mesenchyme and expression of Shh was downregulated in the anterior palatal epithelium in the Bmpr1a conditional mutant embryos, indicating that Bmp signaling regulates mesenchymal-epithelial interactions during palatal outgrowth. In addition, formation of the palatal processes of the maxilla was blocked while formation of the palatal processes of the palatine was significantly delayed, resulting in submucous cleft of the hard palate in the mutant mice. Our data indicate that Bmp signaling plays critical roles in the regulation of palatal mesenchyme condensation and osteoblast differentiation during palatal bone formation.     

Phylogenetic relationships of Asian freshwater Mytiloidea (Bivalvia): a morphological and genetic comparison of Sinomytilus harmandi,Limnoperna fortunei and Septifer bilocularis

ABSTRACT

The Mekong River endemic Sinomytilus harmandi possesses an internal anterior shell septum. It is thus superficially similar to the marine representatives of the, as currently defined, Septiferinae (Mytilidae) although species of the latter possess an anterior adductor muscle located between their septa and are characterised by accessory posterior adductor muscles, both of which are absent in S. harmandi. This study demonstrates that S. harmandi is monophyletic with the only other, similarly native Asian, freshwater mytiloid, Limnoperna fortunei, based on the phylogenetic tree herein presented. Asian freshwaters thus seem to have been colonised by the Mytilidae relatively recently and on two occasions from a common ancestor also associated with representatives of the Lithophaginae, as currently defined. The common presence of an anterior septum in S. harmandi and species of the Septiferinae, as currently recognised, is thus the result of convergence as is the anatomical similarity between S. harmandi and representatives of the Central European Dreissenidae (Heterodonta), which also possess an anterior shell septum.     

Evidence for a frontal-organ homologue in the pineal complex of the salamander,Hynobius dunni

The pineal complex of larval and adult salamanders, Hynobius dunni, was examined by light and scanning electron microscopy. This pineal complex displays an anterior and a posterior portion, both of which possess a lumen. The anterior lumen is small and closed, whereas the posterior lumen is in open communication with the third ventricle. Cell processes of the photoreceptor cells and microvilli of the supportive cells are visible in both lumina. The anterior part of the complex is formed by an independent, second evagination from the common pineal anlage; this process takes place immediately after hatching. The anterior body of the pineal complex of H. dunni appears to be homologous to the frontal organ of anurans.     

Three species of Pycnadenoides Yamaguti, 1938 (Digenea: Opecoelidae), including two new species from temperate marine fishes of Australia

Pycnadenoides pagrosomi Yamaguti, 1938 and P. reversati n. sp. from Pagrus auratus (Sparidae) and P. invenustus n. sp. from Nemadactylus valenciennesi (Cheilodactylidae) are described from the temperate marine waters off south-west Western Australia and south-east Queensland. The difference in the anterior extent of the vitelline follicles observed in P. reversati n. sp. recovered from off south-east Queensland waters and the material from off Western Australia is discussed. P. reversati n. sp. is distinguished from P. pagrosomi mainly in the position of the genital pore and in the arrangement of the testes, and from P. invenustus n. sp. in the posterior extent of the cirrus-sac. P. reversati belongs to the group of species with a short cirrus-sac and P. invenustus to the group with the cirrus-sac reaching into the anterior hindbody.     

Review of the Australian hagfishes with description of two new species of Eptatretus (Myxinidae)

This paper revises and updates taxonomic and distributional information about hagfishes (Myxinidae) from Australia. It covers five species of the genus Eptatretus: Eptatretus cirrhatus known from eastern Australia and also distributed around New Zealand, Eptatretus longipinnis endemic to South Australia, Eptatretus strahani originally described from the Philippines and reported here as a new record from Western Australia and two new species described herein as Eptatretus alastairi and Eptatretus gomoni, both from Western Australia. Eptatretus alastairi is distinguished from all congeners by the unique combination of the following characters: six pairs of gill pouches; three‐cusp multicusps on the anterior and posterior rows of cusps; anterior unicusps 9–12; posterior unicusps 8–11; total cusps 48–56; prebranchial pores 13–16; branchial pores 5–6; trunk pores 50–55; tail pores 11–13; total pores 83–88; two bilaterally symmetrical nasal‐sinus papillae in the dorsal surface of the nasal sinus. Eptatretus gomoni is distinguished from all congeners by the unique combination of the following characters: eight pairs of gill pouches; three‐cusp multicusps on the anterior and two‐cusp multicusps on the posterior row of cusps; anterior unicusps 10–11; posterior unicusps 9–10; total cusps 50; prebranchial pores 12–13; branchial pores 7–8; trunk pores 57–58; tail pores 14–15; total pores 91–93; no nasal‐sinus papillae. An identification key for the Australian species of Eptatretus is also provided.     

Development of the eye-antenna imaginal disc of Drosophila

A clonal analysis of wild-type and aristapedia eye-antenna discs has shown that both discs are subdivided into anterior and posterior compartments. However, the spatial order of the anterior and posterior compartments is reversed in the adult, so that the posterior compartment is at the extreme anterior tip of the fly. The mutation aristapedia transforms both the anterior and the posterior antennal compartments into anterior and posterior leg compartments, respectively. The anteroposterior segregation is established in the eye-antenna disc during the larval period. This contrasts with other discs (leg, wing, proboscis) where the same segregation is established around blastoderm. The engrailed gene is involved in the segregation; some of the mutations in engrailed transform the posterior antennal compartment into a partial mirror image of the anterior one.     

Comparative odontometrical analysis of the first lower molar in <Emphasis Type="Italic">Microtus (Terricola) multiplex</Emphasis> (Arvicolinae,Rodentia) from western Alps

The morphology of the first lower molar (M₁) of Microtus (Terricola) multiplex (Fatio, 1905) was compared amongst 15 populations from the Alps (Switzerland, Italy, France). M. multiplex orientalis from Trentino Alto Adige is close to the nominative subspecies M. multiplex multiplex from Ticino characterised by a great size, a not tilted pitymyan rhombus and an important development of the anterior part of the M₁. M. multiplex druentius from Ubaye mainly differs from the nominative subspecies by a smaller tooth size. Populations from Valle d’Aosta and Piemonte show on the whole a morphology intermediate between M. m. multiplex and M. m. druentius subspecies, however, the pitymyan rhombus is more tilted and the development of the anterior part more reduced in populations from Eastern and Central Piemonte. The Western populations (from Trièvès, Vercors, Royans and Chambaran) belonging to the subspecies M. m. niethammeri are the most differentiated with a small or median size of the M1, a reduced development of the anterior part and a very tilted pitymyan rhombus, particularly in the population from Chambaran. The populations from Matheysine and Grésivaudan are morphologically a link between M. m. druentius and M. m. niethammeri subspecies.     

Growth differentiation factor 11 locally controls anterior–posterior patterning of the axial skeleton

Growth and differentiation factor 11 (GDF11) is a transforming growth factor β family member that has been identified as the central player of anterior–posterior (A–P) axial skeletal patterning. Mice homozygous for Gdf11 deletion exhibit severe anterior homeotic transformations of the vertebrae and craniofacial defects. During early embryogenesis, Gdf11 is expressed predominantly in the primitive streak and tail bud regions, where new mesodermal cells arise. On the basis of this expression pattern of Gdf11 and the phenotype of Gdf11 mutant mice, it has been suggested that GDF11 acts to specify positional identity along the A–P axis either by local changes in levels of signaling as development proceeds or by acting as a morphogen. To further investigate the mechanism of action of GDF11 in the vertebral specification, we used a Cdx2-Cre transgene to generate mosaic mice in which Gdf11 expression is removed in posterior regions including the tail bud, but not in anterior regions. The skeletal analysis revealed that these mosaic mice display patterning defects limited to posterior regions where Gdf11 expression is deficient, whereas displaying normal skeletal phenotype in anterior regions where Gdf11 is normally expressed. Specifically, the mosaic mice exhibited seven true ribs, a pattern observed in wild-type (wt) mice (vs. 10 true ribs in Gdf11^−/− mice), in the anterior axis and nine lumbar vertebrae, a pattern observed in Gdf11 null mice (vs. six lumbar vertebrae in wt mice), in the posterior axis. Our findings suggest that GDF11, rather than globally acting as a morphogen secreted from the tail bud, locally regulates axial vertebral patterning.     

Ichnologic Note A New Ichnospecies of Cardioichnus from the Cretaceous (Albian) of New Mexico

A new ichnospecies of Cardioichnus, Cardioichnus biloba isp. nov., is described and documented from shallow marine strata of Cretaceous (Late Albian) age in southern New Mexico, USA. It is a heart-shaped resting trace which, unlike all other ichnospecies of Cardioichnus, is epichnial and increases in width to a bilobate trace, often with a raised anterior terminus. Only the posterior end of this trace is associated with Bichordites burrows, and the anterior end is bilobate and undisturbed. This represents a resting position which differs from other known species of this cubichnium, which are hypichnial and subquadrate in symmetry. The tracemaker is a spatangoid echinoid, most likely Heteraster d'Orbigny, 1853.