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At least 19 recordsLinked to original sources

Geometric optics of trilobite eyes: a theoretical study of the shape of the aspherical interface in the cornea of schizochroal eyes of phacopid trilobites.

A general method is given to determine theoretically the shape of the aspherical interface that eliminates spherical aberration in an optionally shaped thick lens. The theory is applied to trilobite eyes. On the basis of the geometric optical method presented, the refractive indices and focal length of the original corneal lenses of trilobites can be determined. The shape of the aspherical interface in the cornea of some phacopid trilobites with schizochroal eyes is investigated. The theoretical aspherical interfaces agree well with the real ones.

Animals↗

Zebrafish trilobite identifies new roles for Strabismus in gastrulation and neuronal movements.

Embryonic morphogenesis is driven by a suite of cell behaviours, including coordinated shape changes, cellular rearrangements and individual cell migrations, whose molecular determinants are largely unknown. In the zebrafish, Dani rerio, trilobite mutant embryos have defects in gastrulation movements and posterior migration of hindbrain neurons. Here, we have used positional cloning to demonstrate that trilobite mutations disrupt the transmembrane protein Strabismus (Stbm)/Van Gogh (Vang), previously associated with planar cell polarity (PCP) in Drosophila melanogaster, and PCP and canonical Wnt/beta-catenin signalling in vertebrates. Our genetic and molecular analyses argue that during gastrulation, trilobite interacts with the PCP pathway without affecting canonical Wnt signalling. Furthermore, trilobite may regulate neuronal migration independently of PCP molecules. We show that trilobite mediates polarization of distinct movement behaviours. During gastrulation convergence and extension movements, trilobite regulates mediolateral cell polarity underlying effective intercalation and directed dorsal migration at increasing velocities. In the hindbrain, trilobite controls effective migration of branchiomotor neurons towards posterior rhombomeres. Mosaic analyses show trilobite functions cell-autonomously and non-autonomously in gastrulae and the hindbrain. We propose Trilobite/Stbm mediates cellular interactions that confer directionality on distinct movements during vertebrate embryogenesis.

Animals↗

Role of the zebrafish trilobite locus in gastrulation movements of convergence and extension.

Convergence and extension are gastrulation movements that participate in the establishment of the vertebrate body plan. Using new methods for quantifying convergence and extension movements of cell groups, we demonstrate that in wild-type embryos, dorsal convergence of lateral cells is initially slow, but speeds up between the end of the gastrula period and early segmentation. Convergence and extension movements of lateral cells in trilobite mutants are normal during the gastrula period but reduced by early segmentation. Morphometric studies revealed that during epiboly wild-type gastrulae become ovoid, whereas trilobite embryos remain rounder. By segmentation, trilobite embryos exhibit shorter, broader embryonic axes. The timing of these morphological defects correlates well with impaired cell movements, suggesting reduced convergence and extension are the main defects underlying the trilobite phenotype. Our gene expression, genetic, and fate mapping analyses show the trilobite mutation affects movements without altering dorsoventral patterning or cell fates. We propose that trilobite function is required for cell properties that promote increased speed of converging cells and extension movements in the dorsal regions of the zebrafish gastrula.

Animals↗

Trilobite body patterning and the evolution of arthropod tagmosis.

Preservation permitting patterns of developmental evolution can be reconstructed within long extinct clades, and the rich fossil record of trilobite ontogeny and phylogeny provides an unparalleled opportunity for doing so. Furthermore, knowledge of Hox gene expression patterns among living arthropods permit inferences about possible Hox gene deployment in trilobites. The trilobite anteroposterior body plan is consistent with recent suggestions that basal euarthropods had a relatively low degree of tagmosis among cephalic limbs, possibly related to overlapping expression domains of cephalic Hox genes. Trilobite trunk segments appeared sequentially at a subterminal generative zone, and were exchanged between regions of fused and freely articulating segments during growth. Homonomous trunk segment shape and gradual size transition were apparently phylogenetically basal conditions and suggest a single trunk tagma. Several derived clades independently evolved functionally distinct tagmata within the trunk, apparently exchanging flexible segment numbers for greater regionally autonomy. The trilobite trunk chronicles how different aspects of arthropod segmentation coevolved as the degree of tagmosis increased.

Animals↗

The stability of thoracic segmentation in trilobites: a case study in developmental and ecological constraints.

The decline in origination rate of new metazoan body plans following the Cambrian radiation has been suggested to reflect developmental canalization in derived taxa, limiting their ability to evolve forms with radically different morphotypes. Segmentation is a fundamental aspect of arthropod body plan, and here we show that a derived trilobite that secondarily converged on a morphotype characteristic of basal members of the clade also reverted to a pattern of segmental variability common among basal trilobites. Hence a secular trend in loss of variability of the trilobite thorax was not due to the evolution of an inviolable developmental constraint. This result challenges the notion of developmental canalization in phylogenetically derived taxa. Rather, early variability in trilobites may be the result of ecological factors that promoted segment-rich thoracic morphotypes during Cambrian time.

Animals↗

Post-cambrian trilobite diversity and evolutionary faunas

A cluster analysis of the stratigraphic distribution of all Ordovician trilobite families, based on a comprehensive taxonomic database, identified two major faunas with disjunct temporal diversity trends. The Ibex Fauna behaved as a cohort, declining through the Ordovician and disappearing at the end-Ordovician mass extinction. In contrast, the Whiterock Fauna radiated rapidly during the Middle Ordovician and gave rise to all post-Ordovician trilobite diversity. Its pattern of diversification matches that of the Paleozoic Evolutionary Fauna; hence, trilobites were active participants in the great Ordovician radiations. Extinction patterns at the end of the Ordovician are related to clade size: Surviving trilobite families show higher genus diversity than extinguished families.

Journal Article↗

The Zebrafish trilobite gene is essential for tangential migration of branchiomotor neurons.

Newborn neurons migrate extensively in the radial and tangential directions to organize the developing vertebrate nervous system. We show here that mutations in zebrafish trilobite (tri) that affect gastrulation-associated cell movements also eliminate tangential migration of motor neurons in the hindbrain. In the wild-type hindbrain, facial (nVII) and glossopharyngeal (nIX) motor neurons are induced in rhombomeres 4 and 6, respectively, and migrate tangentially into r6 and r7 (nVII) and r7 (nIX). In all three tri alleles examined, although normal numbers of motor neurons are induced, nVII motor neurons are found exclusively in r4, and nIX-like motor neurons are found exclusively in r6. The migration of other neuronal and nonneuronal cell types is unaffected in tri mutants. Rhombomere formation and the development of other hindbrain neurons are also unaffected in tri mutants. Furthermore, tangential neuronal migration occurs normally in the gastrulation mutant knypek, indicating that the trilobite neuron phenotype does not arise nonspecifically from aberrant gastrulation-associated movements. We conclude that trilobite function is specifically required for two types of cell migration that occur at different stages of zebrafish development.

Animals↗

Exploring developmental modes in a fossil arthropod: growth and trunk segmentation of the trilobite Aulacopleura konincki.

Trilobites offer the opportunity to explore postembryonic development within the fossil record of arthropod evolution. In contrast to most trilobites, the Silurian proetid Aulacopleura konincki from the Czech Republic exhibits marked variation in the mature number of thoracic segments, with five morphs with 18-22 thoracic segments. The combination of abundant articulated specimens available from a narrow stratigraphic interval and segmental intraspecific variation makes this trilobite singularly useful for studying postembryonic growth and segmentation. Trunk segmentation followed a hemianamorphic pattern, as seen in other arthropods and as characteristic of the Trilobita; during a first anamorphic phase, segments were accreted, while in the subsequent epimorphic phase, segmentation did not proceed further despite continued growth. Size increment during the anamorphic phase was targeted and followed Dyar's rule, a geometric progression typical of many arthropods. We consider alternative hypotheses for the control of the switch from anamorphic to epimorphic phases of development. Our analysis favors a scenario in which the mature number of thoracic segments was determined quite early in development rather than at a late stage in association with a critical size threshold. This study demonstrates that hypotheses concerning developmental pattern and control can be tested in organisms belonging to an extinct clade.

Animals↗

Direct evidence for predation on trilobites in the Cambrian.

A fossil arthropod similar to Fuxianhuia and displaying an exceptionally well-preserved alimentary canal with in situ gut contents is reported from the lower Middle Cambrian (ca. 510 Myr ago) of South China. Three-dimensionally preserved, paired phosphatic nodules, arranged in series along both sides of the gut and containing spherical bacteria, probably represent serial digestive glands. The gut is filled with fragments of the eodiscoid trilobite, Pagetia. The well-developed digestive glands and the fragmentary trilobite remains suggest (i) that the arthropod was a durophagous, possibly selective predator, and (ii) that small trilobites such as eodiscoids were a major food source in Cambrian marine ecosystems. This fossil association augments the small number of previously described associations providing more or less direct evidence of predator-prey relationships in Cambrian epibenthic communities.

Animals↗

Image formation by bifocal lenses in a trilobite eye?

In this work we report on a unique and ancient type of eye, in which the lower surface of the upper calcite lens units possessed an enigmatic central bulge making the dioptric apparatus similar to a bifocal lens. This eye belonged to the trilobite Dalmanitina socialis, which became extinct several hundred million years ago. As far as we know, image formation by bifocal lenses of this kind did/does not occur in any other ancient or modern animal visual system. We suggest that the function of these bifocal lenses may be to enable the trilobite to see simultaneously both very near (e.g. floating food particles and tiny preys) and far (e.g. sea floor, conspecifics, or approaching enemies) in the optical environment through the central and peripheral lens region, respectively. This was the only reasonable function we could find to explain the puzzling lens shape. We admit that it is not clear whether bifocality was necessary for the animal studied. We show that the misleading and accidental resemblance of an erroneous correcting lens surface (designed by René DesCartes in 1637 [DesCartes, R. (1637). Oeuvres de DesCartes. La Géometrie. Livre 2. pp. 134. J. Maire, Leyden] to the correcting interface in the compound Dalmanitina lens may be the reason why the earlier students of the Dalmanitina lens did not recognize its possible bifocality.

Accommodation, Ocular↗

Trilobite eyes and the optics of Des Cartes and Huygens.

The thick lenses in the aggregate eyes of a group of trilobites were double structures designed to eliminate spherical aberration. The shape of the optically correcting interface is in accord with constructions by Des Cartes and Huygens and is dictated by a fundamental law of physics. Trilobites may have evolved such sophisticated eye-lenses to maximise optic neurone response in a dimly lit environment.

Adaptation, Physiological↗

Development of the caudal exoskeleton of the pliomerid trilobite Hintzeia plicamarginis new species.

The later juvenile ontogeny of the caudal plate of the early Ordovician pliomerid trilobite Hintzeia plicamarginis new species likely comprised an initial phase during which the rate of appearance of new segments subterminally exceeded that of segment release into the thorax, a short phase of constant segment numbers, and a later phase during which release occurred but in which no new segments appeared. A distinct terminal region became manifest in the second phase. During the second and third phases growth coefficients for individual segments were about 1.1--1.2 per instar. Although the shapes of segments varied during growth, the pattern of ontogenetic shape change appears to have been broadly similar among segments. This suggests an homonomous trunk segment morphology regardless of thoracic or caudal identity in maturity. These results imply that control of trunk exoskeletal segment appearance and articulation were decoupled in this trilobite, and that the terminal region had a distinct mature morphology. H. plicamarginis is described as a new species.

Animals↗

Functional interactions of genes mediating convergent extension, knypek and trilobite, during the partitioning of the eye primordium in zebrafish.

Vertebrate eye development in the anterior region of the neural plate involves a series of inductive interactions dependent on the underlying prechordal plate and signals from the midline of the neural plate, including Hedgehog. The mechanisms controlling the spatiotemporal expression pattern of hedgehog genes are currently not understood. Cyclopia is observed in trilobite (tri) and knypek (kny) mutants with affected convergent extension of the embryonic axis during gastrulation. Here, we demonstrate that tri mutants show a high frequency of partial or complete cyclopia, kny mutants exhibit cyclopia infrequently, while knym119 trim209 double-mutant embryos have dramatically reduced convergent extension and are completely cyclopic. We analyzed the relationships between the convergent extension defect, the expression of hedgehog and prechordal plate genes, and the formation of cyclopia in knym119 and trim209 mutants. Our results correlate the cyclopia phenotype with the abnormal location of hh-expressing cells with respect to the optic primordium. We show that cyclopia in these mutants is not due to an incompetence of tri and kny cells to respond to Hedgehog signaling. Rather, it is a consequence of exceeding a critical distance (>40-50 micrometer) between hedgehog-expressing cells and the prospective eye field. We hypothesize that at this distance, midline cells are not in an appropriate position to physically separate the eye field and that HH and other signals do not reach the appropriate target cells. Furthermore, tri and kny have overlapping functions in establishing proper alignment of the anterior neural plate and midline cells expressing shh and twhh genes when the partitioning of the eye primordium takes place.

Animals↗

Somites in zebrafish doubly mutant for knypek and trilobite form without internal mesenchymal cells or compaction.

In vertebrates, paraxial mesoderm is partitioned into repeating units called somites. It is thought that the mechanical forces arising from compaction of the presumptive internal cells of prospective somites cause them to detach from the unsegmented presomitic mesoderm [1-3]. To determine how prospective somites physically segregate from each other, we used time-lapse microscopy to analyze the mechanics underlying early somitogenesis in wild-type zebrafish and in the mutants trilobite(m209) (tri), knypek(m119) (kny), and kny;tri, which are defective in convergent extension during gastrulation. Formation of somite boundaries in all of these embryos involved segregation, local alignment, and cell-shape changes of presumptive epitheloid border cells along nascent intersomitic boundaries. Although kny;tri somites formed without convergence of the presomitic mesoderm and were composed of only two cells in their anteroposterior (AP) dimension, they still exhibited AP intrasegmental polarity. Furthermore, morphogenesis of somite boundaries in these embryos proceeded in a manner similar to that in wild-type embryos. Thus, intersomitic boundary formation in zebrafish involves short-range movements of presumptive border cells that do not require mechanical forces generated by internal cells or compaction of the presomitic mesoderm.

Animals↗

Parallel gradualistic evolution of Ordovician trilobites.

There are very few high-resolution studies of the fossil record from which to assess the relative frequency of gradualistic and punctuated evolution. Here I report some of the first detailed evidence of phyletic gradualism in benthic macroinvertebrates, based on a study of approximately 15,000 trilobites from central Wales. Over a period of about three million years, as many as eight lineages underwent a net increase in the number of pygidial ribs, a species-diagnostic character. The end members of most lineages have previously been assigned to different species and, in one case, to different genera. In view of intermediate morphologies and temporary trend reversals, however, practical taxonomic subdivision of each lineage proved impossible. The apparent success of earlier Linnean nomenclature (with its implications of discrete species) could easily have been misinterpreted as evidence of punctuation and stasis, and it is probable that detection of many other gradualistic patterns has been hindered by ready application of binominal taxonomy to fossils.

Animals↗

Olenid trilobites: the oldest known chemoautotrophic symbionts?

Late Cambrian to early Ordovician trilobites, the family Olenidae, were tolerant of oxygen-poor, sulfur-rich sea floor conditions, and a case is made that they were chemoautotrophic symbionts. Olenids were uniquely adapted to this habitat in the Lower Paleozoic, which was widespread in the Late Cambrian over Scandinavia. This life habit explains distinctive aspects of olenid morphology: wide thoraces and large numbers of thoracic segments, thin cuticle and, in some species, degenerate hypostome, and the occasional development of brood pouches. Geochemical and field evidence is consistent with this interpretation. Olenids occupied their specialized habitat for 60 million years until their extinction at the end of the Ordovician.

Adaptation, Physiological↗

Trilobite spines and beetle horns: sexual selection in the Palaeozoic?

Raphiophorid trilobites commonly bore median cephalic protuberances such as spines or bulbs, showing a remarkable variety of form. It is unlikely that their primary function was for protection or in hydrodynamics. A case is made that they were secondary sexual features, by comparison with similar morphological structures developed on rhinoceros beetles and other arthropods. This interpretation is supported by four lines of evidence: their ontogeny, their diversity, the existence of plausible examples of sexual dimorphs in some cases and the fact that they show positive allometry.

Animals↗