Combining embryology and paleontology: origins of the anterior-posterior axis in echinoids

The tendency of the periproct to move outside the apical system occurred in eight independent sea urchins lineages, but only one succeeded. The origin of the anterior-posterior axis in irregular echinoids results from that successful attempt, and is primarily achieved by eccentricity of the periproct. The mechanism by which this occurs is best understood through synthesis of data from embryology, paleontology, and phylogenetics. In irregular echinoids, the complete escape of the periproct from the apical system (exocyclism) is a consequence of changes in the embryological sequence that governs periproctal eccentricity. The adaptative significance of this change is discussed. To cite this article: T. Saucède et al., C. R. Palevol 2 (2003).     

Morphological similarity between echinoid taxa as a result of convergent and parallel evolution

Convergence is observed in groups, which are phylogenetically remote. A flat test is typical for some representatives of the order Cassiduloida, such as Jurassic and Early Cretaceous species of the genera Pygurus and Clypeus, and also many Cenozoic «sand dollars,» i.e., echinoids of the order Clypeasteroida. Both usually inhabit coarse sandy grounds of shallow areas. The superorder Spatangacea includes the so-called Echinocorys life form, which is characterized by an oval test with superficial nonpetaloid or subpetaloid ambulacra, marginal or inframarginal periproct, and absence of fascioles. These are the following genera: Early Cretaceous Corthya (family Collyritidae), Late CretaceousLate Paleocene Echinocorys (family Holas-teridae), Paleocene Isaster, Recent Isopatagus (family Isasteridae), Recent Scrippsechinus (family Palaeotro-pidae), and Recent Urechinus (family Urechinidae). In contrast to the majority of spatangaceans with the burrowing mode of life, these genera dwell on the substrate surface. In the Cenozoic, the monobasal apical system appears in some genera of the order Spatangoida, most genera of the order Cassiduloida, all groups of the orders Clypeasteroida and Oligopygoida, and in the genus Echinoneus (order Holectypoida). The paral-lelism is revealed in groups connected by remote relationships. At the end of the Middle Jurassic (Callovian) and, especially, Late Jurassic, the so-called disasterid echinoids (superorder Spatangacea) show a distinct trend to the loss of contact between ocular plates I and V and apices of the posterior ambulacra with the periproct, which are shifted to the anterior part of the apical system (genera Collyrites, Collyropsis, Cyclolam-pas). At the same time, the peristome of some genera was displaced to the anterior margin of the test, which became bilaterally symmetrical in outline. However, in the Jurassic, all spatangaceans remained disasterid echinoids, i.e., had a disjunct apical system, which can be interpreted as a somewhat “abnormal” state. This trend disappeared only at the beginning of the Cretaceous, when “normal” forms with a joint apical system appeared, that is, the families Holasteridae (genera Eoholaster and Holaster, order Holasteroida) and Toxas-teridae (genus Toxaster, order Spatangoida). Interesting examples of synchronous parallelism are provided by the appearance of meridosternous (diasternal) plastron in two collyritid genera (Tetraromania and Corthya) in the Barremian, whereas in the holasterid genus Holaster, this type of plastron apparently appeared in the Valanginian (heterochronous parallelism). The ethmolitic type of the apical system appeared at the end of the Cretaceous and Paleocene at least in five families: Schizasteridae, Paleopneustidae, Brissidae, Spatangidae, and Loveniidae.     

Loriolella,a key taxon for understanding the early evolution of irregular echinoids

Loriolella Fucini is an enigmatic and poorly understood Early Jurassic sea-urchin that has been interpreted both as a regular and an irregular echinoid. The discovery of new and well-preserved material of the type species L. ludovicii (Meneghini) from the Upper Pliensbachian (Lower Jurassic) of Lombardy, Italy, has clarified the morphology of this genus for the first time. Loriolella resembles regular echinoids in having a large (almost certainly monocyclic) apical disc enclosing the periproct, and large primary interambulacral tubercles and spines. However, its peristome is extremely small with only vestigial buccal notches and auricles, and the ambulacra expand adorally forming cassiduloid-like phyllodes. Cladistic analysis suggests that Loriolella is the earliest known microstomatid irregular echinoid. Its unique mixture of primitive and derived characters is important for interpreting the initial steps by which irregular echinoids arose from diadematoid-like regular ancestors.     

Experimental morphology of coronar growth in regular echinoids

Summary The coronar growth of a cidaroid and an aulodont echinoid are investigated by means of tetracycline labelling. The results are compared with earlier investigations on a stirodont and on a camarodont echinoid in order to evaluate the general features of coronar growth. In all echinoids new coronar plates are added at the apical end of the corona throughout the life cycle. The plates are shifted towards the peristome and they grow peripherically.In cidaroids ambulacral (A-) plates are detached from the firm corona. They are transformed into scales covering the peristomial field. The interambulacral (IA-) plates, however, are partially reabsorbed at the peristomial margin. In this manner the oldest solitary interambulacral plates are lost. The subsequent plates are arranged in pairs. The cidaroids thus show interradial growth even at their peristomial margin. This is unique to echinoids.In non-cidaroids there is a perignathic girdle made up of paired ambulacral auricles with interambulacral ridges in between. In some species the ridge is a solitary element. Therefore interradial growth cannot occur in the peristomial margin. In other species the ridge consists of several elements, but it also grows as a whole. Slight resorption of calcite occurs in places at the peristomial margin. In other places, however, calcite is added onto the peristomial edge. In non-cidaroids, therefore, the widening of the peristome is achieved solely by means of lateral growth in the plates bordering the peristome. The shift of the coronar plates from apicad to orad in noncidaroids is a relative shift only.In all echinoids the coronar plates are arranged in meridional columns. All plates grow up to the peristome. Their growth rates are relatively uniform towards the adambulacral sutures (which run between A- and IA-columns). Their growth rates towards the perradius and the interradius respectively are high in younger plates which are positioned above the ambitus, and decrease rapidly in plates located below the ambitus. Near the peristome the interradiad and perradiad growth rates are always considerably lower than adradiad growth rates. Perradial and interradial growth serve to adjust the plates in size and shape to their respective position in the corona.     

Observations on Opisthonecta henneguyi Fauré-Fremiet: Infraciliature, Argyrome, and Myonemic System

The infraciliature, argyrome, and myonemic system of Opisthonecta henneguyi have been examined after impregnation with the Fernández-Galiano silver impregnation method. The oral infraciliature is formed by one haplokinety and three polykineties while the aboral infraciliature is formed by the trochal band—six kinetosomes wide—and the scopula. The trochal band has three fibrillar systems: orally directed fibers, aboral fibers, and oblique fibers. The argyrome is formed by 90 to 135 fine circular striations, which cover the whole organism with the exception of the peristome. The myonemic system is located in two regions: the oral and the aboral ones. The oral myonemic system is formed by a fibrous ring which surrounds the peristome underneath the spiral of the peristomial infraciliature and by a group of fibers which depart from the peristomial disc. The aboral myonemic system is much simpler, and it is formed by fibers which extend radially from the scopula. Biometrical characterizations of the vegetative cells and living cysts were also made, and our data are compared with the results obtained previously by other authors.     

Ontogeny of the holothurian larval nervous system: evolution of larval forms

Echinoderm larvae share numerous features of neuroanatomy. However, there are substantial differences in specific aspects of neural structure and ontogeny between the dipleurula-like larvae of asteroids and the pluteus larvae of echinoids. To help identify apomorphic features, we have examined the ontogeny of the dipleurula-like auricularia larva of the sea cucumber, Holothuria atra. Neural precursors arise in the apical ectoderm of gastrulae and appear to originate in bilateral clusters of cells. The cells differentiate without extensive migration, and they align with the developing ciliary bands and begin neurogenesis. Neurites project along the ciliary bands and do not appear to extend beneath either the oral or aboral epidermis. Apical serotonergic cells are associated with the preoral loops of the ciliary bands and do not form a substantial commissure. Paired, tripartite connectives form on either side of the larval mouth that connect the pre-oral, post-oral, and lateral ciliary bands. Holothurian larvae share with hemichordates and bipinnariae a similar organization of the apical organ, suggesting that the more highly structured apical organ of the pluteus is a derived feature. However, the auricularia larva shares with the pluteus larva of echinoids several features of neural ontogeny. Both have a bilateral origin of neural precursors in ectoderm adjacent to presumptive ciliary bands, and the presumptive neurons move only a few cell diameters before undergoing neurogenesis. The development of the holothurian nervous systems suggests that the extensive migration of neural precursors in asteroids is a derived feature. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Externally visible fluorochrome marks and allometries of growing sea urchins

Externally visible, growing calcitic structures can be marked using fluorochromes. Such marks are useful for field recapture studies in ecology, evolution, and aquaculture as well as for studies on mechanisms of growth and development. We marked 2‐month‐old sea urchins (Strongylocentrotus droebachiensis) with the fluorochromes calcein, calcein blue, and tetracycline by batch‐marking via immersion. Neither growth nor survival was affected by marking. Marks were externally visible on the skeletal plates, demipyramids, and spines of 100% of the marked sea urchins up to 63 d post‐marking. After 342 d, marks were still externally visible on 100% of calcein‐marked, 98% of calcein blue‐marked, and 22% of tetracycline‐marked sea urchins. Marks were brightest on calcein‐marked and faintest on tetracycline‐marked sea urchins, in correspondence to the fluorochrome dose. Growth marks in the aboral oculogenital ring, followed for 333 d, showed that genital plate growth in the hoop direction was greatest adjacent to the anal suture and that both the oculogenital ring and periproct grew less than isometrically. Internal marks (not externally visible) were subsequently seen on 99% of the demipyramids at 342 d post‐marking. Such fluorochrome marks on demipyramids have previously been used to measure aboral and oral‐end demipyramid growth to allometrically calibrate diametrical growth rates of field sea urchins. We found that although aboral demipyramid marks were always clear, 13% of marks on the oral end were obscured. However, we show that measuring only aboral end growth is sufficient for allometric calibration. In a separate experiment, multiple marks of the above three fluorochromes plus alizarin complexone, administered by injection to larger (12.9–37.1 mm diameter) sea urchins, persisted internally for at least 2 years. Multi‐color, internally and externally visible, persistent marks will enhance experimental designs in laboratory, field, and common garden experiments.     

Morphology and Phylogeny of Four New Vorticella Species (Ciliophora: Peritrichia) from Coastal Waters of Southern China

Four new species of Vorticella, V. parachiangi sp. n., V. scapiformis sp. n., V. sphaeroidalis sp. n., and V. paralima sp. n., were isolated from coastal brackish waters of southern China. Their morphology, infraciliature, and silverline system were investigated based on observations of specimens both in vivo and following silver staining. Vorticella parachiangi sp. n. is distinguished by: a J‐shaped macronucleus; a single dorsally located contractile vacuole; a two‐rowed infundibular polykinetid 3, in which row 1 is shorter than row 2; 21–31 silverlines between peristome and aboral trochal band, 6–11 between aboral trochal band and scopula. Vorticella scapiformis sp. n. is characterized by its conspicuously thin and irregularly edged peristomial lip; a J‐shaped macronucleus; a single, ventrally located contractile vacuole; row 1 of the infundibular polykinetid 3 proximally shortened; 18–25 silverlines between peristome and aboral trochal band, 8–12 between aboral trochal band and scopula. Vorticella sphaeroidalis sp. n. can be identified by its small, sub‐spherical zooid; a C‐shaped macronucleus; a ventrally located contractile vacuole; an aboral trochal band adjacent to the scopula; 16–18 silverlines between persitome and aboral trochal band, two between aboral trochal band and scopula. Vorticella paralima sp. n. can be identified by its ovoidal zooid; a J‐shaped macronucleus; a dorsally positioned contractile vacuole; rows 1 and 2 of the infundibular polykinetid 3 proximally shortened; 26–35 silverlines from peristome to aboral trochal band, and 7–13 from aboral trochal band to scopula. The SSU rDNA genes of these four species were sequenced and their phylogeny was analyzed.     

Variabilité architecturale du test chez le genre Collyrites (Echinoidea, Disasteroidea) au Jurassique moyen

The aim of this paper is to study the evolution of the architecture of the « apical system–periproct complex » of the genus Collyrites (Echinoidea, Disasteroidea) for species from the Bathonian and Callovian stages (Middle Jurassic). The studied material comes from several localities of the Paris basin. The apical system of the genus Collyrites is subdivided in two parts: (i) an anterior part composed by four genital (1, 2, 3, 4) and three ocular (II, III, IV) plates, called trivium; (ii) a posterior part composed by two ocular plates (I, V) and one genital plate (5), without gonopor, which circle the periproct, called bivium. The genital plate 5, which may collapse in the periproct, is not always visible. The Bathonian–Callovian transition is marked by a subdivision of the bivium in two parts: the periproct breaks up from the posterior ocular plates (I, V). These morphological changes are associated with architectural modifications. The trivium stays relatively stable during the Bathonian and the Callovian, but a supplementary plate may be inserted into the trivium. The bivium shows important modifications linked to the separation of the periproct and the ocular plates (I and V). Such a separation is marked by a strong development of supplementary plates, these ones keeping in connection with the periproct and the ocular plates I and V. The supplementary plates are more and more developed whereas the distance between the periproct and ocular plates increases. The connection between the trivium and the bivium is similarly provided by supplementary plates. The size of these plates seems to significantly increase between the Bathonian and the Callovian. Moreover, some specimens from the Bathonian and the Callovian may have an atypical architecture with a supplementary genital plate or a genital plate with two pores. The “extraxial axial theory” allows to recognize two types of skeleton: (i) an “axial skeleton” corresponding to ocular plates and plates of the ambulacra and interambulacra; (ii) an “extraxial skeleton” corresponding to the genital and supplementary plates, and the periproct. Architectural modifications between the Bathonian and the Callovian is a result of a more important development of the “extraxial skeleton” while the “axial skeleton” shows few modifications during this time interval.     

Shallow Traces (Pits) in the Test of the Irregular Echinoid Echinocorys scutata Leske from the Chalk (Upper Cretaceous) of the United Kingdom

Specimens of epifaunal irregular echinoids in the Upper Cretaceous of northern Europe have been reported with patterns of circular, nonpenetrative parabolic pits, Oichnus paraboloides (Bromley), in the apical region. Specimens of Echinocorys scutata Leske from the Chalk at two sites in southeast England were commonly penetrated by this trace, generated by an indeterminate pit-forming organism. Pits commonly surround the apical system and, less commonly, occur within it; they occur preferentially anteriorly. Pits occur within plates, not along margins or sutures. Crosscutting of pits indicates that multiple spatfalls probably occurred. The host echinoid added new test plates adjacent to the apical system; thus, plates bearing O. paraboloides were moved abapically. The reduction in number of pits away from the apex, including those with echinoid tubercles reestablished on the base, indicates that, following death of an infester, the echinoid “reclaimed” and infilled them with calcite. The pit-former was most probably an unmineralized invertebrate that used E. scutata as a domicile which provided good access to food-rich currents for suspension feeding. Although the systematic position of the pit-former is unknown, similar infestations are known from other Upper Cretaceous echinoids and Mississippian crinoids.     

Deuterostome evolution: early development in the enteropneust hemichordate, Ptychodera flava

SUMMARY Molecular and morphological comparisons indicate that the Echinodermata and Hemichordata represent closely related sister‐phyla within the Deuterostomia. Much less is known about the development of the hemichordates compared to other deuterostomes. For the first time, cell lineage analyses have been carried out for an indirect‐developing representative of the enteropneust hemichordates, Pty‐ chodera flava. Single blastomeres were iontophoretically labeled with DiI at the 2‐ through 16‐cell stages, and their fates followed through development to the tornaria larval stage. The early cleavage pattern of P. flava is similar to that of the direct‐developing hemichordate, Saccoglossus kowalevskii, as well as that displayed by indirect‐developing echinoids. The 16‐celled embryo contains eight animal “mesomeres,” four slightly larger “macromeres,” and four somewhat smaller vegetal “micromeres.” The first cleavage plane was not found to bear one specific relationship relative to the larval dorsoventral axis. Although individual blastomeres generate discrete clones of cells, the appearance and exact locations of these clones are variable with respect to the embryonic dorsoventral and bilateral axes. The eight animal mesomeres generate anterior (animal) ectoderm of the larva, which includes the apical organ; however, contributions to the apical organ were found to be variable as only a subset of the animal blastomeres end up contributing to its formation and this varies from embryo to embryo. The macromeres generate posterior larval ectoderm, and the vegetal micromeres form all the internal, endomesodermal tissues. These blastomere contributions are similar to those found during development of the only other hemichordate studied, the direct‐developing enteropneust, S. kowalevskii. Finally, isolated blastomeres prepared at either the two‐ or the four‐cell stage are capable of forming normal‐appearing, miniature tornaria larvae. These findings indicate that the fates of these cells and embryonic dorsoventral axial properties are not committed at these early stages of development. Comparisons with the developmental programs of other deuterostome phyla allow one to speculate on the conservation of some key developmental events/mechanisms and propose basal character states shared by the ancestor of echinoderms and hemichordates.     

Development of the nervous system in the brittle star Amphipholis kochii

There are several studies of neural development in various echinoderms, but few on ophiuroids, which develop indirectly via the production of pluteus larvae, as do echinoids. To determine the extent of similarity of neuroanatomy and neural development in the ophiuroids with other echinoderm larvae, we investigated the development of the nervous system in the brittle star Amphipholis kochii (Echinodermata: Ophiuroidea) by immunohistochemistry. Immunoreactive cells first appeared bilaterally in the animal pole at the late gastrula stage, and there was little migration of the neural precursors during A. kochii ontogeny, as is also the case in echinoids and holothuroids. On the other hand, neural specification in the presumptive ciliary band near the base of the arms does occur in ophiuroid larvae and is a feature they share with echinoids and ophiuroids. The ophiopluteus larval nervous system is similar to that of auricularia larvae on the whole, including the lack of a fine network of neurites in the epidermis and the presence of neural connections across the oral epidermis. Ophioplutei possess a pair of bilateral apical organs that differ from those of echinoid echinoplutei in terms of relative position. They also possess coiled cilia, which may possess a sensory function, but in the same location as the serotonergic apical ganglia. These coiled cilia are thought to be a derived structure in pluteus-like larvae. Our results suggest that the neural specification in the animal plate in ophiuroids, holothuroids, and echinoids is a plesiomorphic feature of the Ambulacraria, whereas neural specification at the base of the larval arms may be a more derived state restricted to pluteus-like larvae.     

Ultrastructure of tentacles in the sipunculid worm <Emphasis Type="Italic">Thysanocardia nigra</Emphasis> Ikeda, 1904 (Sipuncula)

The ultrastructure of the tentacles was studied in the sipunculid worm Thysanocardia nigra. Flexible digitate tentacles are arranged into the dorsal and ventral tentacular crowns at the anterior end of the introvert of Th. nigra. The tentacle bears oral, lateral, and aboral rows of cilia; on the oral side, there is a longitudinal groove. Each tentacle contains two oral tentacular canals and an aboral tentacular canal. The oral side of the tentacle is covered by a simple columnar epithelium, which contains large glandular cells that secrete their products onto the apical surface of the epithelium. The lateral and aboral epithelia are composed of cuboidal and flattened cells. The tentacular canals are lined with a flattened coelomic epithelium that consists of podocytes with their processes and multiciliated cells. The tentacular canals are continuous with the radial coelomic canals of the head and constitute the terminal parts of the tentacular coelom, which shows a highly complex morphology. Five tentacular nerves and circular and longitudinal muscle bands lie in the connective tissue of the tentacle wall. Similarities and differences in the tentacle morphology between Th. nigra and other sipunculan species are discussed.Original Russian Text Copyright © 2005 by Biologiya Morya, Maiorova, Adrianov.     

The nature and construction of skeletal spines inPocillopora damicornis (Linnaeus)

Extreme variability in the size, shape and spacing of skeletal spines ofPocillopora damicornis has been demonstrated both within single colonies and also between colonies from different environments. Preliminary studies indicated that the majority of spines from branch tips at the apex of the colony display a ‘fasciculate’ growth surface in contrast to partly fasciculate or ‘smooth’ growth surfaces exhibited by spines from branch tips at the base of the colony. No significant differences in the height and width of costal spines from apical and basal branch tips within a single colony were observed, although spines from colonies exposed to strong wave action tended to be significantly shorter and narrower than those from more sheltered environments. Both costal and coenosteal spines from wave-exposed colonies displayed branching and divided extremities while those from sheltered environments consisted of simple cones. Spines develop as an outgrowing of the calicoblastic ectoderm which secretes the skeleton. Growing costal and coenosteal spines are enveloped by a layer of calicoblastic ectoderm which penetrates through mesogloea, aboral gastroderm, coelenteron, oral gastroderm, mesogloea and finally oral ectoderm. Spines within the corallite are surrounded by calicoblastic ectoderm, mesogloea and aboral gastroderm only. A scheme for the growth of the spines is discussed.     

Localization of sensory mechanisms utilized by coral planulae to detect settlement cues

Coral planulae are induced to settle and metamorphose by contact with either crustose coralline algae or marine bacterial biofilms. Larvae of two coral species, Pocillopora damicornis and Montipora capitata, which respond to different metamorphic cues, were utilized to investigate the sensory mechanisms used to detect metamorphic cues. Because the aboral pole of the coral planula is the point of attachment to the substratum, we predicted that it is also the point of detection for cues. To determine where sensory cells for cues are localized along the body, individual larvae were transversely cut into oral and aboral portions at various levels along the oral–aboral axis, and exposed to settlement‐inducing substrata. Aboral ends of M. capitata metamorphosed, while oral ends continued to swim. However, in larvae of P. damicornis, ¾ oral ends (i.e., lacking the aboral pole) were also able to metamorphose, indicating that the cells that detect cues may be distributed along the sides of the body. These cells do not correspond to FMRFamide‐immunoreactive cells that are present throughout the body. Cesium ions induced both aboral and oral ends of larvae of both species to settle, suggesting that oral ends have not lost their capacity to metamorphose, despite lacking sensory cells to detect natural cues. To determine whether sensory cells in larvae of P. damicornis are restricted to one side of the body, swimming behavior over substrata was observed in larvae labeled with diI, a red fluorescent lipophilic membrane stain. The larvae were found to rotate around the oral–aboral axis, with their surface against the substratum, not favoring a particular side for detecting cues. While clarifying the regions of the larval body important for settlement and metamorphosis in coral planulae, we conclude that significant differences between coral species may be due to differences in the distribution of sensory structures in relation to different planular sizes.     

Cyathotheca suecica and its bearing on the evolution of the Edrioasteroidea

The European Ordovician edrioasteroid Cyathotheca has a variable tubular theca composed of a single plate. The oral surface is covered by five large inter-radial deltoids fused to biserial ambulaeral cover plates, and a periproct in the C-D inter-radius accompanied by two or three adanal plates. Cyathotheca lacks ambulaeral flooring plates. The American and Baltic Ordovician genus Cyathocystis is identical except that it has a complete ring of small peripheral plates surrounding the deltoids. Both genera are assigned to the family Cyathocystidae. The Devonian genera Timeischytes and Hadrochthus are regarded as neotenous offshoots from the Agelacrinitidae and unrelated to cyathocystids. True cyathocystids arose neotenously from the Cambrian stromatocystitids by attachment of the centro-dorsal and failure to develop further aboral or ambulaeral plates. This branch represents a separate evolutionary line to the main stromatocystitid-edrioasterid-isorophid line.     

SYMMETRY,LOCOMOTION, AND THE EVOLUTION OF AN ANTERIOR END: A LESSON FROM SEA URCHINS

Bilaterally symmetrical, “regular” sea urchins in the Family Echinometridae (Class Echinoidea; Phylum Echinodermata) were found to lack a locomotor anterior. Heterocentrotus mammillatus and Echinometra mathaei were observed while locomoting. Members of both ellipsoidal species were found to proceed with their short or long axis foremost with statistically equivalent frequencies. This finding demonstrates that the evolution of bilateral symmetry is not always accompanied by the evolution of a locomotor “anterior” end. The elliptical echinometrid sea urchins provide a particularly appropriate study group for investigating the relationship between the evolution of body form and locomotor behavior. Although the radially symmetrical regular sea urchins, from which the echinometrids sprang, lack a locomotor anterior, all “irregular” echinoids, which are also derived from a regular ancestor but are bilaterally symmetrical, possess an “obligate” locomotor anterior. The symmetry and behavior exhibited by the elliptical echinometrid sea urchins therefore demonstrates that the first irregular echinoids (which exhibit bilateral symmetry by definition) need not have possessed a locomotor anterior as they do today.     

Small bending,big curvature

<正>The classical "Cholodny-Went theory" predicted that directional stimuli trigger the redistribution of auxin, which governs the differential growth of plant organs through potent effects on cell expansion, thereby establishing an"auxin-then-growth" paradigm; this theory has been validated for both gravitropism and phototropism in plants(reviewed in Muthert et al., 2020).     

Comparative microanatomy of the branchial sieve in three sympatric cyprinid species,related to filter-feeding mechanisms

The chimaeroid holocephalian fishes are distinguished among extant chondrichthyans by the possession of three pairs of tooth plates, evergrowing and partially hypermineralized, that are not shed and replaced like the teeth of living elasmobranchs. Although derivation of the chimaeroid tooth plate from the fusion of members of a plesiomorphic chondrichthyan tooth family has been proposed, evidence for this hypothesis has been lacking. A new analysis of the development and structure of the tooth plates in Callorhinchus milii (Holocephali, Chimaeriformes) reveals the compound nature of the tooth plates in a chimaeroid fish. Each tooth plate consists of an oral and aboral territory that form independently in the embryo and maintain separate growth surfaces through life. The descending lamina on the aboral surface of the tooth plate demarcates the growth surface of the aboral territory. Comparison with the tooth plates of Chimaera monstrosa indicates that compound tooth plates may be a feature of all chimaeroids in which a descending lamina is present. The tooth plates in these fishes represent the fusion of two members of a reduced tooth family. The condition of the tooth plates in C. milii is plesiomorphic for chimaeroids and is of evolutionary significance in that it provides further evidence to support a lyodont dentition in chimaeroid fishes similar to that found in other chondrichthyans. © 1994 Wiley-Liss, Inc.     

Ultrastructure of the type “B” cells in the rectal pad epithelium of Locusta migratoria

The electron microscopical structure of the type “B” cells in the rectal pad epithelium of Locusta is described. The type “B” cells occur singly in the distal region of the rectal pad epithelium. They are characteristically goblet shaped and join with contiguous type “A” or rectal pad cells, near the apical surface by means of a restricted region of septate desmosomes. Type “B” cells possess a microvillate apical membrane, with the villi arranged as a rosette overlying the apical inaginations of adjacent type “A” cells. Large numbers of microtubules and vacuoles of various sizes containing an assortment of inclusions are present in the apical region of the type “B” cells. Many of the microtubules insert distally on hemidesmosomes located in the apical plasma membrane. Rough endoplasmic reticulum and mitochondria are also present but neither are abundant. The possible significance of these findings is discussed.