Search

Oviparous amniotes produce a large yolky egg that gives rise to a free-living hatchling. Structural characteristics and functional attributes of the egg are best known for birds, which have a large mass of fluid yolk surrounded by an extraembryonic yolk sac. Yolk nutrients are delivered to the embryo via the vascular yolk sac. This developmental pattern and nutrient transport mechanism is thought to be representative of all other lineages of amniotes. Recent discovery of a snake with cellularized yolk organized around a meshwork of blood vessels reveals an additional pattern for yolk mobilization, which may also occur in other squamate reptiles (lizards and snakes). This complex yolk sac raises interesting questions about developmental mechanisms and suggests a possible model for the transition between the egg of anamniotes and that of amniotes.

Although the development, anatomy, and physiology of the vertebrate olfactory system are fairly well understood, there is still no clear definition of the terminal nerve complex acknowledged by all. Among the most debated matters is whether or not the extrabulbar projections found in anamniotes should or should not be considered part of the terminal nerve complex. In this context, we investigated the early development of the extrabulbar pathway in Xenopus larvae from placodal differentiation to postmetamorphic stages. We showed that the extrabulbar fibers become visible around Stage 42 and are conserved throughout metamorphosis. We confirmed previous reports concerning their central projection patterns. In addition, we showed that these fibers originate from two types of cell bodies located in the olfactory epithelium at premetamorphic stages. Furthermore, in postmetamorphic animals, we showed that the extrabulbar axons originated from both aquatic and aerial cavities. Retrograde tracing experiment also revealed densifications evocating cell bodies along the extrabulbar axons, distributed at different positions along the olfactory nerve depending on the stages of development. These densifications were observed closer to the periphery early in development and always closer to the olfactory bulb up to the metamorphic climax. We discuss these results in light of the latest theories and more recent reports. Anat Rec, 296:1462-1476, 2013. © 2013 Wiley Periodicals, Inc.

Classical hypotheses regarding the evolutionary origin of paired appendages propose transformation of precursor structures (gill arches and lateral fin folds) into paired fins. During development, gnathostome paired appendages form as outgrowths of body wall somatopleure, a tissue composed of somatic lateral plate mesoderm (LPM) and overlying ectoderm. In amniotes, LPM contributes connective tissue to abaxial musculature and forms ventrolateral dermis of the interlimb body wall. The phylogenetic distribution of this character is uncertain because lineage analyses of LPM have not been generated in anamniotes. We focus on the evolutionary history of the somatopleure to gain insight into the tissue context in which paired fins first appeared. Lampreys diverged from other vertebrates before the acquisition of paired fins and provide a model for investigating the preappendicular condition. We present vital dye fate maps that suggest the somatopleure is eliminated in lamprey as the LPM is separated from the ectoderm and sequestered to the coelomic linings during myotome extension. We also examine the distribution of postcranial mesoderm in catshark and axolotl. In contrast to lamprey, our findings support an LPM contribution to the trunk body wall of these taxa, which is similar to published data for amniotes. Collectively, these data lead us to hypothesize that a persistent somatopleure in the lateral body wall is a gnathostome synapomorphy, and the redistribution of LPM was a key step in generating the novel developmental module that ultimately produced paired fins. These embryological criteria can refocus arguments on paired fin origins and generate hypotheses testable by comparative studies on the source, sequence, and extent of genetic redeployment.

In the mature mammalian and avian central nervous systems, neuronal destructions are followed by reactive gliosis, but data on other vertebrates are rather controversial. Mammals and birds belong to different amniote groups (Synapsida and Diapsida, respectively), but exhibit common general features in their glial architecture, mainly the predominance of astrocytes. Two vertebrate groups seem to be in special positions of glial evolution: turtles (Testudiniformes) and skates and rays (Batoidea). The purely ependymoglial system of turtles seems to be the simplest one among the extant amniotes. In skates and rays, true astrocytes are preponderant glial elements, in contrast to the other "anamniotes" (and even to reptiles). We investigated stab wounds by the immunohistochemical detection of GFAP in turtles (Trachemys-formerly Pseudemys-scripta elegans), a skate (Raja clavata) and rays (Dasyatis akajei and Torpedo marmorata). Sharks (Scyliorhinus canicula) as ependymoglia-predominated chondrichthyans, and-for positive controls-rats were also studied. In the elasmobranchs, other astroglial markers: glutamine synthetase and S100 protein were also applied. Neither turtles nor elasmobranchs presented considerable astroglial reactions. Critically surveying the former reports on different vertebrates, these results complete the picture that typical post-lesion reactive gliosis is confined to mammals and birds. Analysis of the astroglial systems from phylogenetic perspective suggests that the capability of forming glial demarcation and scar formation evolved independently in mammals and birds. Predominance of astrocytes is a necessary condition but not sufficient for reactive gliosis. The intense glial reactivity of mammals and birds may be attributed to their complex cerebralization.

In the present chapter we have analyzed comparative aspects of the organization of the diencephalon and hypothalamus in nonmammalian vertebrates. We focused primarily on the neuroanatomical features highlighted by developmental patterns of gene expression, which are strikingly conserved throughout all vertebrate classes and that are readily compared across species when analyzed using the current paradigm provided by the prosomeric model for the organization of the forebrain.

Reptiles are the unique ectothermic amniotes, providing the key link between ectothermic anamniotes fish and amphibians, and endothermic birds and mammals; becoming an important group to study with the aim of providing significant knowledge into the evolutionary history of vertebrate immunity. Classification systems for reptiles' leukocytes have been described by their appearance rather than function, being still inconsistent. With the advent of modern techniques and the establishment of analytical protocols for snakes' blood by flow cytometry, we bring a qualitative and quantitative assessment of innate activities presented by snakes' peripheral blood leukocytes, thereby linking flow cytometric features with fluorescent and light microscopy images. Moreover, since corticosterone is an important immunomodulator in reptiles, hormone levels of all blood samples were measured. We provide novel and additional information which should contribute to better understanding of the development of the immune system of reptiles and vertebrates.

The complexity of the pallium during evolution has increased dramatically in many different respects. The highest level of complexity is found in mammals, where most of the pallium (cortex) shows a layered organization and neurons are generated during development following an inside-out order, a sequence not observed in other amniotes (birds and reptiles). Species-differences may be related to major neurogenetic events, from the neural progenitors that divide and produce all pallial cells. In mammals, two main types of precursors have been described, primary precursor cells in the ventricular zone (vz; also called radial glial cells or apical progenitors) and secondary precursor cells (called basal or intermediate progenitors) separated from the ventricle surface. Previous studies suggested that pallial neurogenetic cells, and especially the intermediate progenitors, evolved independently in mammalian and sauropsid lineages. In the present study, we examined pallial neurogenesis in the amphibian Xenopus laevis, a representative species of the only group of tetrapods that are anamniotes. The pattern of pallial proliferation during embryonic and larval development was studied, together with a multiple immunohistochemical analysis of putative progenitor cells. We found that there are two phases of progenitor divisions in the developing pallium that, following the radial unit concept from the ventricle to the mantle, finally result in an outside-in order of mature neurons, what seems to be the primitive condition of vertebrates. Gene expressions of key transcription factors that characterize radial glial cells in the vz were demonstrated in Xenopus. In addition, although mitotic cells were corroborated outside the vz, the expression pattern of markers for intermediate progenitors differed from mammals.

Sensory hair cell losses lead to hearing and balance deficits that are permanent for mammals, but temporary for nonmammals because supporting cells in their ears give rise to replacement hair cells. In mice and humans, vestibular supporting cells grow exceptionally large circumferential F-actin belts and their junctions express E-cadherin in patterns that strongly correlate with postnatal declines in regeneration capacity. In contrast, chicken supporting cells retain thin F-actin belts throughout life and express little E-cadherin. To determine whether the junctions in chicken ears might be representative of other ears that also regenerate hair cells, we investigated inner ears from dogfish sharks, zebrafish, bullfrogs, Xenopus, turtles, and the lizard, Anolis. As in chickens, the supporting cells in adult zebrafish, Xenopus, and turtle ears retained thin circumferential F-actin belts and expressed little E-cadherin. Supporting cells in adult sharks and bullfrogs also retained thin belts, but were not tested for E-cadherin. Supporting cells in adult Anolis exhibited wide, but porous webs of F-actin and strong E-cadherin expression. Anolis supporting cells also showed some cell cycle reentry when cultured. The results reveal that the association between thin F-actin belts and low E-cadherin is shared by supporting cells in anamniotes, turtles, and birds, which all can regenerate hair cells. Divergent junctional specializations in supporting cells appear to have arisen independently in Anolis and mammals. The presence of webs of F-actin at the junctions in Anolis appears compatible with supporting cell proliferation, but the solid reinforcement of the F-actin belts in mammals is associated with its absence. J. Comp. Neurol., 521:1430–1448, 2013. © 2012 Wiley Periodicals, Inc.

Neuronal production in metazoans is tightly controlled by Delta/Notch-dependent signals regulating lateral inhibition. It is currently thought that lateral inhibition takes place in clusters of precursors with equal capacity to trigger and receive Notch-dependent inhibitory signals. However, this view neglects crucial dynamical aspects of the process. In this review, we discuss two of these dynamic factors, whose alterations yield dysfunctions in neurogenesis. First, precursors show variable neurogenic capacity as they go through the cell cycle. Second, differentiating precursors are in direct contact with non-neurogenic cells at the wavefront of expanding neurogenic domains. We discuss the mechanisms adopted by Metazoa to prevent these dysfunctions in the lateral inhibitory process, which include cell cycle synchronization occurring in the invertebrate neural epithelium and during primary neurogenesis in anamniotes, interkinetic nuclear movement in the vertebrate neuroepithelium and generalized Delta expression ahead of the neurogenic wavefront. The emerging concept is that lateral inhibition during neurogenesis occurs in dynamic clusters of precursors and requires specific mechanisms to avoid distortions resulting from the interaction between neurogenic and non-neurogenic precursors. The advance in visualizing Notch dynamics with real-time imaging at cellular and subcellular levels will notably contribute to our understanding of these novel "aspects of motion" in neurogenesis.

Neural circuit development is an activity-dependent process. This activity can be spontaneous, such as the retinal waves that course across the mammalian embryonic retina, or it can be sensory-driven, such as the activation of retinal ganglion cells (RGCs) by visual stimuli. Whichever the source, neural activity provides essential instruction to the developing circuit. Indeed, experimentally altering activity has been shown to impact circuit development and function in many different ways and in many different model systems. In this review, we contemplate the idea that retinal waves in amniotes, the animals that develop either in ovo or utero (namely reptiles, birds and mammals) could be an evolutionary adaptation to life on land, and that the anamniotes, animals whose development is entirely external (namely the aquatic amphibians and fish), do not display retinal waves, most likely because they simply don’t need them. We then review what is known about the function of both retinal waves and visual stimuli on their respective downstream targets, and predict that the experience-dependent development of the tadpole visual system is a blueprint of what will be found in future studies of the effects of spontaneous retinal waves on instructing development of retinorecipient targets such as the superior colliculus (SC) and the lateral geniculate nucleus.

FGIN-1-27 is an agonist at the translocator protein 18 kDa (TSPO), a cholesterol transporter that is associated with neurosteroidogenesis. This protein has been identified as a peripheral binding site for benzodiazepines; in anamniotes, however, a second TSPO isoform that is absent in amniotes has been implicated in erythropoiesis. Functional conservation of the central benzodiazepine-binding site located in the GABAA receptors has been demonstrated in anamniotes and amniotes alike; however, it was not previously demonstrated for TSPO. The present investigation explored the behavioral effects of FGIN-1-27 on an anxiety test in zebrafish (Danio rerio, Family: Cyprinide) and on a mixed anxiety/panic test on wall lizards (Tropidurus oreadicus, Family: Tropiduridae). Results showed that FGIN-1-27 reduced anxiety-like behavior in the zebrafish light/dark preference test similar to diazepam, but with fewer sedative effects. Similarly, FGIN-1-27 also reduced anxiety- and fear-like behaviors in the defense test battery in wall lizards, again producing fewer sedative-like effects than diazepam; the benzodiazepine was also unable to reduce fear-like behaviors in this species. These results A) underline the functional conservation of TSPO in defensive behavior in anamniotes; B) strengthen the proposal of using anamniote behavior as models in behavioral pharmacology; and C) suggest TSPO/neurosteroidogenesis as a target in treating anxiety disorders.

Vertebrate neck musculature spans the transition zone between head and trunk. The extent to which the cucullaris muscle is a cranial muscle allied with the gill levators of anamniotes or is instead a trunk muscle is an ongoing debate. Novel computed tomography datasets reveal broad conservation of the cucullaris in gnathostomes, including coelacanth and caecilian, two sarcopterygians previously thought to lack it. In chicken, lateral plate mesoderm (LPM) adjacent to occipital somites is a recently identified embryonic source of cervical musculature. We fate-map this mesoderm in the axolotl (Ambystoma mexicanum), which retains external gills, and demonstrate its contribution to posterior gill-levator muscles and the cucullaris. Accordingly, LPM adjacent to the occipital somites should be regarded as posterior cranial mesoderm. The axial position of the head-trunk border in axolotl is congruent between LPM and somitic mesoderm, unlike in chicken and possibly other amniotes. DOI: http://dx.doi.org/10.7554/eLife.09972.001, eLife digest Muscles in the head and trunk (main body) form from different parts of the embryo, and their development uses different genes. Trunk muscles are derived from somites – paired blocks of cells arranged in segments on either side of the midline (which divides the body into left and right halves). By contrast, cells that give rise to head muscles are arranged in a continuous mass. But what about neck muscles? Some studies claim they develop like head muscles; others suggest they are trunk muscles. These studies commonly examine mice or chickens. By examining species that have a more primitive complement of head and neck muscles, Sefton et al. now show that a neck muscle should be considered a kind of head muscle. Gill muscles are definitive head muscles. Sefton et al. found that the cucullaris, a prominent neck muscle in fishes and amphibians, forms from the same mass of cells that gives rise to gill muscles. Moreover, studying muscle development in Mexican axolotls showed that cells that contribute to gill muscles extend into the trunk, which is further back in the embryo than was previously known. Previous studies reported the cucullaris muscle is absent in a “lobe-finned” fish called the coelacanth, which is closely related to four-limbed animals. However, by using a technique called micro-computed tomography to visualize the neck muscles of this fish, Sefton et al. show that the cucullaris muscle is present and connects the rear-most gill to the shoulder. The finding that neck muscles form like head muscles in the axolotl confirms a previous claim that was based on studies of bird embryos. A future challenge is to understand the molecular and genetic mechanisms that establish the boundary between head and trunk muscles, and work out how those mechanisms might have influenced how the neck evolved. DOI: http://dx.doi.org/10.7554/eLife.09972.002

The axial skeleton is a defining feature of vertebrates and is patterned during somitogenesis. Cyclically expressed members of the notch and other signaling pathways, described as the ‘segmentation clock’, regulate the formation of somite boundaries. Comparisons among vertebrate model systems have revealed fundamental shifts in the regulation of expression among critical genes in the notch pathway. However, insights into the evolution of these expression differences have been limited by the lack of information from non-avian reptiles. We analyzed the segmentation clock of the first Lepidosaurian reptile sequenced, the green anole lizard, Anolis carolinensis, for comparison with avian and mammalian models. Using genomic sequence, RNA-Seq transcriptomic data, and in situ hybridization analysis of somite-stage embryos, we carried out comparative analyses of key genes and found that the anole segmentation clock displays features common to both amniote and anamniote vertebrates. Shared features with anamniotes, represented by Xenopus laevis and Danio rerio, include an absence of lunatic fringe (lfng) expression within the presomitic mesoderm (PSM), a hes6a gradient in the PSM not observed in the chicken or mouse, and EGF repeat structure of the divergent notch ligand, dll3. The anole and mouse share cycling expression of dll1 ligand in the PSM. To gain insight from an Archosaurian reptile, we analysed LFNG and DLL1 expressions in the American alligator. LFNG expression was absent in the alligator PSM, like the anole but unlike the chicken. In contrast, DLL1 expression does not cycle in the PSM of the alligator, similar to the chicken but unlike the anole. Thus, our analysis yields novel insights into features of the segmentation clock that are evolutionarily basal to amniotes versus those that are specific to mammals, Lepidosaurian reptiles, or Archosaurian reptiles.

Networked gene activities control the evolutionarily conserved histogenetic organization of the central nervous system of vertebrates. Genoarchitectonic studies contribute to the analysis of each morphogenetic field by identifying distinct progenitor domains and corresponding derivatives whose pattern of gene expression shows a unique combinatory code. Previous studies in the pretectal region (caudal diencephalon) have defined three anteroposterior genoarchitectonic domains that are conserved in birds and mammals. Here, we have studied the embryonic pretectal genoarchitecture in the amphibian Xenopus laevis, in order to determine whether it is possible to define a comparable anteroposterior tripartition of the amphibian pretectal area. The expression patterns of 14 genes mapped from early embryonic stages to metamorphic climax allowed us to define the boundaries of the pretectum, the expected precommissural, juxtacommissural, and commissural anteroposterior domains, and some dorsoventral subdivisions. Taken together, our data provide evidence for a conserved pattern of pretectal domains and subdomains, shared by amniotes and amphibian anamniotes (tetrapods), understandable as part of a general Bauplan in vertebrates. J. Comp. Neurol. 519:1024‐1050, 2011.

The testes of Hemidactylus turcicus are composed of seminiferous tubules lined with continuous germinal epithelia in which multiple germ cell morphologies can be found during the active months of sperm development. Spermatogenesis is quiescent during September, with only spermatogonia A and spermatogonia B present in the seminiferous epithelia and minimal mitotic activity is observed. Recrudescence begins in October and the early stages of spermatogenesis progress through November. The onset of spermiation is observed in December and continues through August with the heaviest sperm release occurring in June and July. Multiple generations of late elongated spermatids are found in association with early mitotic and meiotic cells during the months of December–August. This temporal germ cell development strategy is similar to that described in other squamates and anamniotes and is different from the spatial development exhibited by birds and mammals, in which germ cell populations collectively progre...

Ground Skink (Scincella lateralis) testes were examined histologically to determine the testicular organization and germ cell development strategy employed during spermatogenesis. Testicular tissues were collected from 19 ground skinks from Aiken County, South Carolina during the months of March-June, August, and October. The testes consisted of seminiferous tubules lined with germinal epithelia in which germ cells matured in close association with Sertoli cells. As germ cells matured, they migrated away from the basal lamina of the epithelia towards the lumina of the seminiferous tubules. The testes were spermatogenically active during the months of March, April, May, June, and October (largest seminiferous tubule diameters and epithelial heights), but entered a quiescent period in August (smallest seminiferous tubule diameter and epithelial height) where only spermatogonia type A and B and early spermatocytes were present in low numbers within the seminiferous epithelium. Although the testicular organization was similar to other amniotes, a temporal germ cell development strategy was employed during spermatogenesis within Ground Skinks, similar to that of anamniotes. Thus, this skink's germ cell development strategy, which also has been recently reported in all other major reptilian clades, may represent an evolutionary intermediate in terms of testicular organization between anamniotes and birds and mammals.

Some animals, notably birds, preferentially approach and capture food items in their right visual field. However, this lateralised behaviour has not been studied extensively in anamniotes. Here we test eye preference during feeding for a fish, (Perccottus glenii; Teleostei, Perciformes), a newt, (Pleurodeles walti; Amphibia, Caudata), and a frog, (Xenopus laevis; Amphibia, Anura) using a test chamber that assesses reaction to visual stimuli while blocking olfactory and mechanical input. Both the fish and the newt showed right preferences in reactions to food items, but the frog did not. Our data extend our knowledge of the lateralised behaviours of vertebrates and are the first record of lateralised prey capture in a caudate amphibian. This finding dates back the history of the common pattern for visual lateralisation in vertebrates to Devonian, when the fish and quadruped lineages diverged.

In eukaryotes, in contrast to prokaryotes, phyletic lineages are usually quite distinct. Therefore, the best classifications of eukaryotes are usually phylogeny-based. However, in many groups of organisms, higher rank taxa are based on horizontal rather than phyletic groups or, more precisely, on groups of the mixed type. This is largely true for vertebrates where the division into fishes and tetrapods or into anamniotes (nonamniotes) and amniotes is just of the mixed type. Frequent parallelisms in these cases make the strict adherence to the principle of monophyly senseless and imply a compromise system. Strict cladistic practice violates this approach and ignores Hennig’s rule that his methodology should only be applied to synchronous organisms.

During land adaptation of the integument in tetrapods, an efficient stratum corneum was originated through the evolution of numerous corneous proteins in addition to the framework of intermediate filament-keratins present in keratinocytes. The new genes for corneous proteins were originated in a chromosome region indicated as epidermal differentiation complex (EDC), a locus with no apparent relationship to keratin genes. The addition of EDC proteins to IF-keratins transformed the process of epidermal keratinization present in anamniotes into a new process of cornification in the epidermis and skin appendages of amniotes, including hairs and feathers. In sauropsids among other EDC proteins a peculiar type of small proteins evolved a central region of 34 amino acids conformed as beta-sheets that, differently from the other EDC proteins, allowed the formation of long polymers of filamentous proteins customarily termed beta-keratins but in the present review reclassified as EDC corneous beta proteins. To the initial beta-sheets present in the corneous beta proteins specific N- and C-regions were later added in the proteins of different sauropsids in relation to the evolution of the corneous layer and skin appendages. Cornification contributed to the evolutive success of amniotes in the terrestrial environment.

The olfactory and vomeronasal systems of vertebrates are characterised by neurogenesis occurring throughout life. The regenerative ability of olfactory receptor neurons relies on specific glial cells, the olfactory and vomeronasal axon-surrounding cells. Numerous studies have examined mammalian olfactory ensheathing cells which are considered potential candidates for spinal cord injury repair using cell-based therapy. With regard to non-mammalian vertebrates, limited information is available on these glial cells in fish, and there is no information on them in terrestrial anamniotes, the amphibians. In the present research, we studied the immunocytochemical characteristics of axon-surrounding cells in Ambystoma mexicanum. Urodeles have relatively simple olfactory and vomeronasal systems, and represent a good model for studying ensheathing cells in extant representatives of basal tetrapods. Sections from the decalcified heads of A. mexicanum were immunocytochemically processed for the detection of proteins used in research on mammalian olfactory-ensheathing cells. S100, GFAP and NCAM were clearly observed. p75NTR, Gal-1 and PSA-NCAM showed weak staining. No vimentin immunopositivity was observed. The corresponding areas of the olfactory and vomeronasal pathways displayed the same staining characteristics, with the exception of Gal-1, p75NTR and PSA-NCAM in the mucosae. The degree of marker expression was not uniform throughout the sensory pathways. In contrast to fish, both olfactory and vomeronasal nerves displayed uniform staining intensity. This study showed that some markers for mammalian and fish-ensheathing glia are also applicable in urodeles. The olfactory systems of vertebrates show similarities, and also clear dissimilarities. Further investigations are required to ascertain the functional significance of these regional and interspecific differences.

Summary: It has been one and a half centuries since Enrico Sertoli published the seminal discovery of the testicular 'nurse cell', not only a key cell in the testis, but indeed one of the most amazing cells in the vertebrate body. In this review, we begin by examining the three phases of morphological research that have occurred in the study of Sertoli cells, because microscopic anatomy was essentially the only scientific discipline available for about the first 75 years after the discovery. Biochemistry and molecular biology then changed all of biological sciences, including our understanding of the functions of Sertoli cells. Immunology and stem cell biology were not even topics of science in 1865, but they have now become major issues in our appreciation of Sertoli cell's role in spermatogenesis. We end with the universal importance and plasticity of function by comparing Sertoli cells in fish, amphibians, and mammals. In these various classes of vertebrates, Sertoli cells have quite different modes of proliferation and epithelial maintenance, cystic vs. tubular formation, yet accomplish essentially the same function but in strikingly different ways. © 2016 American Society of Andrology and European Academy of Andrology.

The Lower Permian fossiliferous infills of the Dolese Brothers Limestone Quarry, near Richards Spur, Oklahoma, have preserved the most diverse assemblage of Paleozoic terrestrial vertebrates, including small-bodied reptiles and lepospondyl anamniotes. Many of these taxa were previously known only from fragmentary remains, predominantly dentigerous jaw elements and numerous isolated skeletal elements. The recent discovery of articulated skulls and skeletons of small reptiles permits the recognition that dentigerous elements, previously assigned at this locality to the anamniote lepospondyl Euryodus primus, belong to a new captorhinid eureptile, Opisthodontosaurus carrolli gen. et sp. nov. This mistaken identity points to a dramatic level of convergence in mandibular and dental anatomy in two distantly related and disparate clades of terrestrial tetrapods and sheds light on the earliest instance of durophagy in eureptiles.

The correct formation of all central nervous system tissues depends on the proper balance of neurogenesis and developmental cell death. A model system for studying these programs is the ganglion cell layer (GCL) of the vertebrate retina because of its simple and well-described structure and amenability to experimental manipulations. The GCL contains approximately equal numbers of ganglion cells and displaced amacrine cells. Ganglion cells are the first or among the first cells born in the retina in all the studied vertebrates. Neurogenesis and cell death have been studied extensively in the GCL of various amniotes (rodents, chicks, and monkeys) and anamniotes (fish and frogs), and the two processes highlight developmental differences between the groups. In amniotes, neurogenesis occurs during a defined period prior to birth/hatch or the opening of the eyes, whereas in anamniotes, neurogenesis extends past hatching into adulthood—sometimes for years. Roughly half of GCL neurons die during development in amniotes, whereas developmental cell death does not occur in the GCL neurons of anamniotes. This review discusses the spatial and temporal patterns of neurogenesis, cell death, and possible explanation of cell death in the GCL. It also examines markers widely used to distinguish between ganglion cells and displaced amacrine cells, and methods employed to birth date neurons.

Research over the last few years has demonstrated that the amygdaloid complex in amniotes shares basic developmental, hodological and neurochemical features. Furthermore, homolog territories of all main amygdaloid subdivisions have been recognized among amniotes, primarily highlighted by the common expression patterns for numerous developmental genes. With the achievement of new technical approaches, the study of the precise neuroanatomy of the telencephalon of the anuran amphibians has been possible, revealing that most of the structures present in amniotes are recognizable in these anamniotes. Thus, recent investigations have yielded enough results to support the notion that the organization of the anuran amygdaloid complex includes subdivisions with origin in ventral pallial and subpallial territories, a strong relationship with the vomeronasal and olfactory systems, abundant intra-amygdaloid connections, a main output center involved in the autonomic system, profuse amygdaloid fiber systems, and distinct chemoarchitecture. When all these new data about the development, connectivity and neurochemistry of the amygdaloid complex in anurans are taken into account, it becomes patent that a basic organization pattern is shared by both amniotic and anamniotic tetrapods.

Gribbins, K.M., Happ, C.S. and Sever, D.M. 2005. Ultrastructure of the reproductive system of the black swamp snake (Seminatrix pygaea). V. The temporal germ cell development strategy of the testis. — Acta Zoologica (Stockholm) 86: 223-230 The germ cell development strategy during spermatogenesis was investigated in the black swamp snake (Seminatrix pygaea). Testicular tissues were collected, embedded in plastic, sectioned by ultramicrotome, and stained with methylene blue and basic fuchsin. Black swamp snakes have a postnuptial pattern of development, where spermatogenesis occurs from May to July and spermiation is completed by October. Though spatial relationships are seen between germ cells within the seminiferous epithelium during specific months, accumulation of spermatogonia and spermatocytes early in spermatogenesis and the depletion of spermatocytes and accumulation of spermatids late in spermatogenesis prevent consistent cellular associations. This temporal germ cell development within an amniotic testis is consistent with that seen in other recently studied temperate reptiles (slider turtle and wall lizard). These reptiles' temporal development is more similar to the developmental strategy found in anamniotes than the spatial germ cell development that characterizes birds and mammals. Our findings also imply that a third germ cell development strategy may exist in temperate breeding reptiles. Because of the phylogenetic position of reptiles between anamniotes and other terrestrial amniotes, this common germ cell development strategy shared by temperate reptiles representing different orders may have significant implications as far as the evolution of sperm development within vertebrates.

In eukaryotes, ribosome synthesis largely takes place in a specialized nuclear domain - the nucleolus. It has recently become apparent that this organelle is involved in the biogenesis of most cellular ribonucleoprotein particles (RNPs), as well as in cell-cycle regulation, making it central to gene expression. The field has traditionally acknowledged that each nucleolus is organized in three morphologically distinct compartments. Here, however, we discuss our view that in fact many eukaryotes have bipartite nucleoli. We propose that, during evolution, a third nucleolar compartment emerged at the transition between the anamniotes and the amniotes, following a substantial increase in size of the rDNA intergenic region. We believe that these conclusions have important implications for understanding the structure-function relationships within this key cellular organelle.

We used two developmental transcription factors, x-Eomes (T-box family) and x-Lhx5 (LIM-homeodomain family), to follow the origin and development of the olfactory bulbs in Xenopus. During embryonic and larval development, x-Eomes and x-Lhx5 were expressed in highly similar patterns, in the lateral and latero-ventral wall of the pallium. In adults, both markers were strongly and specifically expressed in mitral cells, i.e., in the projection neurons of the main and accessory olfactory bulbs. These results demonstrate the pallial origin of the olfactory projecting cells in Xenopus. Combined with previous results suggesting a subpallial origin for olfactory interneurons, these findings emphasize the dual origin of different neuronal populations in the bulbs of anamniotes, and suggest that this organization is a shared feature of tetrapods.

Adult fish and salamanders regenerate specific neurons as well as entire CNS areas after injury. Recent studies shed light on how these anamniotes activate progenitor cells, generate the required cell types, and functionally integrate these into a complex environment. Some developmental signals and mechanisms are recapitulated during neuronal regeneration, whereas others are unique to the regeneration process. The use of genetic techniques, such as cell ablation and lineage-tracing, in combination with cell-type-specific expression profiling reveal factors that initiate, fine-tune, and terminate the regenerative response in anamniotes, with a view to translating findings to non-regenerating species.

The hippocampus in humans and other mammals is essential for episodic and relational memories. Comparative evidence indicates that a hippocampal pallium homologue is present in birds, reptiles, amphibians, ray-finned fishes, cartilaginous fishes and agnathans. Some of their characteristics, such as the topological position and the pattern of connectivity, appear remarkably well conserved. We review here substantial data showing that in all the vertebrate groups studied up to date, from fish to mammals, the hippocampus plays a fundamental role in spatial memory. In these vertebrates groups, the hippocampal pallium homologue is involved in the use of map-like, relational representations of the objective space that provide stable allocentric frames of reference, thus allowing flexible navigation. These similarities suggest a common evolutionary ancestry and indicate that the functional properties of the hippocampus appear early in the vertebrate phylogenesis and are retained through the independent evolution of the vertebrate lineages. Ministerio de Economía y Competitividad de España PSI2011-27489

The basic architecture of the mammalian neocortex is remarkably similar across species. Pallial structures in the reptilian brain are considered amniote precursors of mammalian neocortex, whereas pallia of anamniotes ("lower" vertebrates) have been deemed largely insignificant with respect to homology. Here, we examine the cytoarchitecture of the lateral pallium in the lamprey, the phylogenetically oldest group of extant vertebrates. We reveal a three-layered structure with similar excitatory cell types as in the mammalian cortex and GABAergic interneurons. The ventral parts are sensory areas receiving monosynaptic thalamic input that can be activated from the optic nerve, whereas the dorsal parts contain motor areas with efferent projections to the brainstem, receiving oligosynaptic thalamic input. Both regions receive monosynaptic olfactory input. This three-layered "primordial" lamprey lateral pallium has evolved most features of the three-layered reptilian cortices and is thereby a precursor of the six-layered "neo" cortex with a long-standing evolutionary precedent (some 500 million years ago).

A comparative analysis of catecholaminergic systems in the brain and spinal cord of vertebrates forces to reconsider several aspects of the organization of catecholamine systems. Evidence has been provided for the existence of extensive, putatively catecholaminergic cell groups in the spinal cord, the pretectum, the habenular region, and cortical and subcortical telencephalic areas. Moreover, putatively dopamine- and noradrenaline-accumulating cells have been demonstrated in the hypothalamic periventricular organ of almost every non-mammalian vertebrate studied. In contrast with the classical idea that the evolution of catecholamine systems is marked by an increase in complexity going from anamniotes to amniotes, it is now evident that the brains of anamniotes contain catecholaminergic cell groups, of which the counterparts in amniotes have lost the capacity to produce catecholamines. Moreover, a segmental approach in studying the organization of catecholaminergic systems is advocated. Such an approach has recently led to the conclusion that the chemoarchitecture and connections of the basal ganglia of anamniote and amniote tetrapods are largely comparable. This review has also brought together data about the distribution of receptors and catecholaminergic fibers as well as data about developmental aspects. From these data it has become clear that there is a good match between catecholaminergic fibers and receptors, but, at many places, volume transmission seems to play an important role. Finally, although the available data are still limited, striking differences are observed in the spatiotemporal sequence of appearance of catecholaminergic cell groups, in particular those in the retina and olfactory bulb.

We analyzed the medaka optic tectum (OT) morphogenesis by using 5-bromo-2’deoxyuridine (BrdU) immunohistochemistry (with a new method we developed for pulselabeling embryos) and in situ hybridization with three probes, two for recently cloned homeobox genes (Ol-Prx3 [Paired-Related-Homeobox3] and Ol-Gsh1 [Genetic-ScreenHomeobox1]) and one for Ol-tailless. The tectal anlage first appears as a sheet of proliferating cells expressing Ol-Gsh1 and Ol-tailless but not Ol-Prx3. Cells subsequently cease to proliferate in a superficial and rostral zone and begin to express Ol-Prx3. When tectal lamination begins, the proliferative zone (mpz) becomes restricted to a crescent at the OT medial, caudal, and lateral margin. This mpz functions throughout the fish’s entire life. It produces cells that are added at the OT’s edge as radial rows, spanning every layer of the OT. The cells of the mpz continue to express Ol-tailless in the adult, whereas Ol-Gsh1 expression is turned off. When superficial layers form, Ol-Prx3 expression becomes restricted to the underlying deep layer, where it persists in the adult. Ol-Prx3 seems to be a marker for the differentiation of a subset of deep cells and allows analysis of tectal lamination, whereas Ol-tailless and Ol-Gsh1 could be involved in the control of tectal cell proliferation. This study constitutes a first step toward molecular approach to OT development in anamniotes. We compare and discuss the expression patterns of the homologs of the genes studied, and more generally the morphogenetic patterns of the medaka tectum, with those encountered in other cortical structures and in other vertebrate groups. J. Comp. Neurol. 413:385‐404, 1999. r 1999 Wiley-Liss, Inc.

Some anamniotic aquatic vertebrates lay eggs in a terrestrial habitat that is hostile to the survival of hatchings or larvae. These terrestrial eggs are ready and able to hatch at a particular developmental time, but do not hatch until presented with suitable conditions for aquatic larval survival. Beyond this time, hatching is possible whenever aquatic conditions occur. The duration of extended terrestrial incubation is dependent on the availability of energy for metabolism from the yolk. Extended incubation is useful for anamniotic eggs laid in terrestrial habitats where conditions suitable for larval survival arrive with unpredictable or variable timing. Examples of anamniotes with delayed hatching and extended terrestrial incubation can be found among teleost fishes, anurans, and caudate amphibians. This paper characterizes the embryonic period, compares this mode with other forms of developmental plasticity in anamniotes, evaluates the constraints and advantages of this life history mode, and examines how some fishes and amphibians are able to obtain the benefits of terrestriality for their eggs when the timing of the return to aquatic conditions is not entirely predictable.

Genome size and GC-percent were determined by means of a special method of DNA flow cytometry in 154 vertebrate species. For the total dataset, a highly significant positive correlation was found between both parameters. The overall distribution of points is not linear but triangular: a wide range of GC-percent values is observed at the lower end of genome size range, whereas with an increase in genome size the lower limit for GC-percent is elevated, gradually approaching the upper limit (about 46%). In teleost fishes, which occupy the lower part of genome size range, the negative relationship between both parameters was observed. Two positive linear relationships were found between mean genome size and GC-percent of the main vertebrate groups (one includes fishes, amphibians, and mammals, the other consists of reptiles and birds, which show the higher GC-percent for their genome sizes). Distribution of variance between taxonomic levels indicates that GC-percent is more evolutionarily conservative than genome size in anamniotes. Anuran amphibians show the greatest part of genome size variability at the lower taxonomic levels as compared to other vertebrates (with no additional variance already above the genus level). The data obtained with different methods are compared. It is shown that the proposed method can provide useful data for studies on genome evolution and biodiversity.

The segmental series of somites in the vertebrate embryo gives rise to the axial skeleton. In amniote models, single vertebrae are derived from the sclerotome of two adjacent somites. This process, known as resegmentation, is well-studied using the quail-chick chimeric system, but the presumed generality of resegmentation across vertebrates remains poorly evaluated. Resegmentation has been questioned in anamniotes, given that the sclerotome is much smaller and lacks obvious differentiation between cranial and caudal portions. Here, we provide the first experimental evidence that resegmentation does occur in a species of amphibian. Fate mapping of individual somites in the Mexican axolotl (Ambystoma mexicanum) revealed that individual vertebrae receive cells from two adjacent somites as in the chicken. These findings suggest that large size and segmentation of the sclerotome into distinct cranial and caudal portions are not requirements for resegmentation. Our results, in addition to those for zebrafish, indicate that resegmentation is a general process in building the vertebral column in vertebrates, although it may be achieved in different ways in different groups.

Because our knowledge of cholinergic systems in the brains of amphibians is limited, the present study aimed to provide detailed information on the distribution of cholinergic cell bodies and fibers as revealed by immunohistochemistry with antibodies directed against the enzyme choline acetyltransferase (ChAT). To determine general and derived features of the cholinergic systems within the class of Amphibia, both anuran (Rana perezi, Xenopus laevis) and urodele (Pleurodeles waltl) amphibians were studied. Distinct groups of ChAT-immunoreactive cell bodies were observed in the basal telencephalon, hypothalamus, habenula, isthmic nucleus, isthmic reticular formation, cranial nerve motor nuclei, and spinal cord. Prominent plexuses of cholinergic fibers were found in the olfactory bulb, pallium, basal telencephalon, ventral thalamus, tectum, and nucleus interpeduncularis. Comparison of these results with those obtained in other vertebrates, including a segmental approach to correlate cell populations, reveals that the cholinergic systems in amphibians share many features with amniotes. Thus, cholinergic pedunculopontine and laterodorsal tegmental nuclei could be identified in the amphibian brain. The finding of weakly immunoreactive cells in the striatum of Rana, which is in contrast with the condition found in Xenopus, Pleurodeles, and other anamniotes studied so far, has revived the notion that basal ganglia organization is more preserved during evolution than previously thought. J. Comp. Neurol. 382:499-534, 1997. © 1997 Wiley-Liss Inc.

In common with the embryos of other anamniotes, young of the Australian lungfish, Neoceratodus forsteri, have ciliated cells in the epidermis. These first appear at stage 28, ˜ 10 days before hatching, and develop progressively to a peak in numbers and in activity at stage 44, just after hatching. After this point, ciliary action in the epidermal cells slowly declines, and cilia disappear completely from the outer surface of the hatchling by stage 52. Cilia are lost earlier from the oral epithelium, between stages 45 and 46, and from the epithelium covering the gills and lining the operculum at stage 51, although they are retained in the nares and in the cavity of the olfactory organ. To assess possible functions for the ciliated epidermis in lungfish hatchlings, the presence of cilia in the epidermis of young N. forsteri is compared with landmarks of development. The ciliated epidermal cells are not associated with movements of the embryo within the egg capsule, nor are they a part of a feeding mechanism. They are not related to oxygen uptake. The ciliated epidermis appears to function as a mechanism for clearing the animal of particles and settling organisms before hatching, when the egg membranes have developed holes, and after hatching, when the young fish is living among the submerged rootlets of trees growing on the river bank or in dense stands of aquatic plants. The function of a ciliated epidermis in N. forsteri hatchlings in relation to microhabitat is discussed. © 1996 Wiley-Liss, Inc.

Previous studies have shown that Agr genes, which encode thioredoxin domain-containing secreted proteins, play a critical role in limb regeneration in salamanders. To determine the evolutionary conservation of Agr function, it is important to examine whether Agrs play a similar role in species with a different type of regeneration. Here, we refined the phylogeny of Agrs, revealing three subfamilies: Ag1, Agr2 and Agr3. Importantly, we established that Ag1 was lost in higher vertebrates, which correlates with their decreased regeneration ability. In Xenopus laevis tadpoles (anamniotes), which have all three Agr subfamilies and a high regenerating capacity, Agrs were activated in the stumps of tails and hindlimb buds that were amputated at stage 52. However, Agrs were not up-regulated when the hindlimb buds were amputated at stage 57, the stage at which their regeneration capacity is lost. Our findings indicate the general importance of Agrs for body appendages regeneration in amphibians.

Background: The axial skeleton is one of the defining evolutionary landmarks of vertebrates. How this structure develops and how it has evolved in the different vertebrate lineages is, however, a matter of debate. Vertebrae and vertebral structures are derived from the embryonic somites, although the mechanisms of development are different between lineages. Discussion: Using the anecdotal description of a teratological newt (Triturus dobrogicus) with an unusual malformation in its axial skeleton, we review, compare, and discuss the development of vertebral structures and, in particular, the development of centra from somitic cellular domains in different vertebrate groups. Vertebrae development through re-segmentation of the somitic sclerotomal cells is considered the general mechanism among vertebrates, which has been generalized from studies in amniotic model organisms. The prevalence of this mechanism among anamniotes is, however, controversial. We propose alternative developmental mechanisms for vertebrae formation that should be experimentally tested. Summary: Research in model organisms, especially amniotes, is laying the foundations for a thorough understanding of the mechanisms of development of the axial skeleton in vertebrates, foundations that should expand the extent of future comparative studies. Although immersed in the ‘-omics’ era, we emphasize the need for an integrative and organismal approach in evolutionary developmental biology for a better understanding of the causal role of development in the evolution of morphological diversity in nature., JV was supported by the NKFP project (Faunagenezis 3B-02304) from the National Office for Research and Technology, Hungary, and the Hungarian Scientific Research Fund (OTKA K84071). DB was partially supported by the National Science Foundation EF-0334939 project (USA) and a JAE-DOC fellowship from the CSIC (Spain) under the program “Junta para la Ampliación de Estudios” co-financed by the European Social Fund (ESF). The authors also thank the Unit of Information Resources for Research (URICI-CSIC) for the co-financing of this publication in Open Access., We also acknowledge institutional support from the Unit of Information Resources for Research at the “Consejo Superior de Investigaciones Científicas” (CSIC) for the article-processing charges contribution., SI

The retina of anamniotes (fish and amphibia), unlike the CNS of most vertebrates, can regenerate neurons following injury. Using the highly ordered mosaic of single and double cones in the retina of the adult green sunfish (Lepomis cyanellus) as our model system, we examined the events that followed the surgical excision of a small patch of central retina. After surgery there was a transient elevation in the number, and a change in the distribution, of proliferative cells within the retina. The wound was filled in two ways: a proliferative regeneration of new retina and a nonproliferative movement of the wound boundaries toward the center of the lesion. The nonproliferative movement stretched the surrounding, intact retina. In stretched retina the basic pattern of the cone mosaic was maintained, but it was augmented by new cones, even though cones are not normally generated in intact central retina. The stretch itself likely triggered the anomalous cone production. The new and preexisting cones in stretched retina had their morphological phenotypes influenced by mutual contact, often resulting in atypical morphologies (triple and quadruple cones). In the center of the lesioned area, the regenerated cone mosaic was disordered, had a higher than normal cone density, and contained atypical morphologies. The presence of outer segments and synaptic pedicles suggested that the new cones in regenerated and stretched retina were functional. We interpret these results to mean (1) a stretch-induced decrease in cell density can trigger a compensatory, adaptive neurogenesis, (2) cone morphological phenotypes in fish retina are plastic throughout life, and are influenced by cone-cone contacts, (3) the mechanisms that spatially regulate cone production during normal growth are disrupted regeneration.

The distribution of calretinin-like immunopositive cells and fibers in the optic tectum of the tench (Tinca tinca) was studied by using a polyclonal antibody and the avidin-biotin-peroxidase technique. A clear laminated pattern of calretinin-like immunoreactivity was observed. The stratum periventriculare demonstrated a large number of strongly labeled cells whereas in the strata album centrale and griseum centrale, and at the boundary between the strata griseum centrale and fibrosum et griseum superficiale, some scarce, weakly immunostained cells were observed. No immunoreactive cells were seen in the strata fibrosum et griseum superficiale, opticum and marginale. Cells belonging to neuronal types X and XIV, previously characterized using Golgi impregnation, were found to be calretinin-like immunoreactive. Most calretinin-like immunopositive fibers were found in the strata fibrosum et griseum superficiale and opticum with a distribution pattern similar to retinotectal axons in these layers. In agreement with previous biochemical studies, our data suggest that, by contrast to all other classes of vertebrates, instead of calretinin and calbindin D-28k, only one protein is present in teleosts. Nevertheless, the calretinin-like immunostaining pattern in the teleost optic tectum was more complex than that previously described for calbindin D-28k. When compared to the calretinin-immunostaining in the rat superior colliculus, it is evident the presence in both amniotes and anamniotes of calretinin-immunopositive retinotectal axons. However, the distribution patterns of intrinsic calretinin-immunoreactive cells were different.(ABSTRACT TRUNCATED AT 250 WORDS)

Publisher Summary Neuroanatomical concepts of the hypothalamus, the forebrain territory that controls homeostasis and drives in vertebrates, are over 100 years old. Neuroanatomists of the 19th century generally considered the hypothalamus to consist of the tuberal or infundibular area, a median prominence at the brain ventral surface at the location of the pituitary gland, framed in front and at the sides by the optic chiasm and tracts. The neural plate fate maps of amniotes and anamniotes are strictly comparable from a topological point of view. The relative positions of the prospective brain areas can be extrapolated from one species to another. The prospective longitudinal or axial dimension of the brain is best represented by the neural/non-neural border of the neural plate, which will transform into the longitudinal roof plate of the closed neural tube. This developmental analysis suggests that all adult median forebrain derivatives found between the anterior commissure and the mamillary area are equally rostralmost loci in the brain. At the open neural plate, the rostral forebrain primordium contains the prospective hypothalamus as a large area centered upon the alar, basal, and floor portions of the terminal wall, its caudal part covering as well a rostral part of the lateral wall, whereas the prospective telencephalon forms a thin band at the dorsal periphery of the alar portion of these domains, including the corresponding roof area. A handful of genes, coding either for transcription factors or secreted protein morphogens, participate in early patterning and regional identity specification along the AP and DV dimensions of the whole forebrain, including the hypothalamus. The complex networked interplay of many molecular signals that control brain development represents a dynamic ontogenetic system that tends to reach different equilibrium states in different parts of the neuroepithelial wall.

The morphogenesis of vertebrate body parts remains an open question. It is not clear whether the existence of different structures, such as a head, can be addressed by fundamental laws of tissue movement and deformation, or whether they are only a sequence of stop-and-go genetic instructions. I have filmed by time-lapse microscopy the formation of the presumptive head territory in chicken embryos. I show that the early lateral evagination of the eye cups and of the mesencephalic plate is a consequence of a sudden change in boundary conditions of the initial cell flow occurring in these embryos. Due to tissue flow, and collision of the two halves of the embryo, the tissue sheet movement is first dipolar, and next quadrupolar. In vivo air puff tonometry reveals a simple visco-elastic behaviour of the living material. The jump from a dipolar to a quadrupolar flow changes the topology of the early morphogenetic field which is observed towards a complex vortex winding with a trail (the eye cups and brain folds). The hydrodynamical model accounts for the discontinuity of the vector field at the moment of collision of the left and right halves of the embryo, at a quantitative level. This suggests a possible mechanism for the morphogenesis of the head of amniotes, as compared to cephalochordates and anamniotes.

Major common features have been reported for the organization of the basal telencephalon in amniotes, and most characteristics were thought to be acquired in the transition from anamniotes to amniotes. However, gene expression, neurochemical, and hodological data obtained for the basal ganglia and septal and amygdaloid complexes in amphibians (anamniotic tetrapods) have strengthened the idea of a conserved organization in tetrapods. A poorly characterized region in the forebrain of amniotes has been the bed nucleus of the stria terminalis (BST), but numerous recent investigations have characterized it as a member of the extended amygdala. Our study analyzes the main features of the BST in anuran amphibians to establish putative homologies with amniotes. Gene expression patterns during development identified the anuran BST as a subpallial, nonstriatal territory. The BST shows Nkx2.1 and Lhx7 expression and contains an Islet1-positive cell subpopulation derived from the lateral ganglionic eminence. Immunohistochemistry for diverse peptides and neurotransmitters revealed that the distinct chemoarchitecture of the BST is strongly conserved among tetrapods. In vitro tracing techniques with dextran amines revealed important connections between the BST and the central and medial amygdala, septal territories, medial pallium, preoptic area, lateral hypothalamus, thalamus, and prethalamus. The BST receives dopaminergic projections from the ventral tegmental area and is connected with the laterodorsal tegmental nucleus and the rostral raphe in the brainstem. All these data suggest that the anuran BST shares many features with its counterpart in amniotes and belongs to a basal continuum, likely controlling similar reflexes, reponses, and behaviors in tetrapods. J. Comp. Neurol. 520:330–363, 2012. © 2011 Wiley Periodicals, Inc.

Synopsis Reptiles are a karyologically heterogeneous group, where some orders and suborders exhibit characteristics similar to those of anamniotes and others share similarities with homeotherms. The class also shows different evolutionary trends, for instance in genome and chromosome size and composition. The turtle DNA base composition is similar to that of mammals, whereas that of lizards and snakes is more similar to that of anamniotes. The major karyological differences between turtles and squamates are the size and composition of the genome and the rate at which chromosomes change. Turtles have larger and more variable genome sizes, and a greater amount of middle repetitive DNA that differs even among related species. In lizards and snakes size of the genome are smaller, single-copy DNA is constant within each suborder, and differences in repetitive DNA involve fractions that become increasingly heterogeneous with widening phylogenetic distance. With regard to variation in karyotype morphology, turtles and crocodiles show low variability in chromosome number, morphology, and G-banding pattern. Greater variability is found among squamates, which have a similar degree of karyotypic change—as do some mammals, such as carnivores and bats—and in which there are also differences among congeneric species. An interesting relationship has been highlighted in the entire class Reptilia between rates of change in chromosomes, number of living species, and rate of extinction. However, different situations obtain in turtles and crocodiles on the one hand, and squamates on the other. In the former, the rate of change in chromosomes is lower and the various evolutionary steps do not seem to have entailed marked chromosomal variation, whereas squamates have a higher rate of change in chromosomes clearly related to the number of living species, and chromosomal variation seems to have played an important role in the evolution of several taxa. The different evolutionary trends in chromosomes observed between turtles and crocodiles on the one hand and squamates on the other might depend on their different patterns of G-banding.

This review summarizes vertebrate rhombic lip and early cerebellar development covering classic approaches up to modern developmental genetics which identifies the relevant differential gene expression domains and their progeny. Most of this information is derived from amniotes. However, progress in anamniotes, particularly in the zebrafish, has recently been made. The current picture suggests that rhombic lip and cerebellar development in jawed vertebrates (gnathostomes) share many characteristics. Regarding cerebellar development, these include a ptf1a expressing ventral cerebellar proliferation (VCP) giving rise to Purkinje cells and other inhibitory cerebellar cell types, and an atoh1 expressing upper rhombic lip giving rise to an external granular layer (EGL, i.e., excitatory granule cells) and an early ventral migration into the anterior rhombencephalon (cholinergic nuclei). As for the lower rhombic lip (LRL), gnathostome commonalities likely include the formation of precerebellar nuclei (mossy fiber origins) and partially primary auditory nuclei (likely convergently evolved) from the atoh1 expressing dorsal zone. The fate of the ptf1a expressing ventral LRL zone which gives rise to (excitatory cells of) the inferior olive (climbing fiber origin) and (inhibitory cells of ) cochlear nuclei in amniotes, has not been determined in anamniotes. Special for the zebrafish in comparison to amniotes is the predominant origin of anamniote excitatory deep cerebellar nuclei homologues (i.e., eurydendroid cells) from ptf1a expressing VCP cells, the sequential activity of various atoh1 paralogues and the incomplete coverage of the subpial cerebellar plate with proliferative EGL cells. Nevertheless, the conclusion that a rhombic lip and its major derivatives evolved with gnathostome vertebrates only and are thus not an ancestral craniate character complex is supported by the absence of a cerebellum (and likely absence of its afferent and efferent nuclei) in jawless fishes.

Until recently, the histology and ultrastructural events of spermatogenesis in reptiles were relatively unknown. Most of the available morphological information focuses on specific stages of spermatogenesis, spermiogenesis, and/or of the mature spermatozoa. No study to date has provided complete ultrastructural information on the early events of spermatogenesis, proliferation and meiosis in class Reptilia. Furthermore, no comprehensive data set exists that describes the ultrastructure of the entire ontogenic progression of germ cells through the phases of reptilian spermatogenesis (mitosis, meiosis and spermiogenesis). The purpose of this review is to provide an ultrastructural and histological atlas of spermatogenesis in reptiles. The morphological details provided here are the first of their kind and can hopefully provide histological information on spermatogenesis that can be compared to that already known for anamniotes (fish and amphibians), birds and mammals. The data supplied in this review will provide a basic model that can be utilized for the study of sperm development in other reptiles. The use of such an atlas will hopefully stimulate more interest in collecting histological and ultrastructural data sets on spermatogenesis that may play important roles in future nontraditional phylogenetic analyses and histopathological studies in reptiles.

The distribution in the spinal cord of the trigeminal primary projections in the frog Rana ridibunda was studied by means of the anterograde transport of horseradish peroxidase (HRP). Upon entering the medulla via the single trigeminal root, a conspicuous descending tract that reaches the cervical spinal cord segments is established. This projection arises in the ophthalmic (V1), maxillary (V2), and mandibular (V3) trigeminal nerve subdivisions. In the spinal cord, only a minor somatotopic arrangement of the trigeminal fibers was observed, with the fibers arising in V3 terminating somewhat more medially than those from V1 and V2. A dense projection to the medial aspect of the spinal cord, above the central canal, primarily involves V3. Each trigeminal branch sends projections at cervical levels to the contralateral dorsal field, and those from V2 are most abundant. Bilateral experiments with HRP application show convergence of primary trigeminal and spinal afferents within the dorsal field of the spinal cord. The pattern of arrangement of the trigeminal primary afferent fibers in the spinal cord of this frog largely resembles that of amniotes. However, the organization seems simpler and the slight somatotopic distribution of V1, V2, and V3 fibers is similar to the condition in other anamniotes. © 1993 Wiley-Liss, Inc.

This note presents a mechanistic explanation of the transition between the morphology of cephalochordates to that of amniotes. By a careful study of the morphogenetic movements which occur during the early stages of development of a typical amniote (a chicken embryo), we are able to show that the formation of a vertebrate body follows a sequence: first, formation of dorsal folds, then head and heart as dorsal and ventral folds, and finally another dorsal fold, which eventually builds up the chorion. This order has a physical origin linked to the velocity field of the tissue flow. These folds form at right angles to the flow direction, and the topology of the chordates flow is hyperbolic. This mechanism explains the differences between the successive bauplans, by the cumulate forward and backward movement of the flow. Eventually, the entire phenomenon can be described as a self-organized system of Russian dolls, by which the heart finds itself inside the embryo, and the embryo itself inside the chorion. In addition, the phenomenon has a mirror symmetry in the anterior and in the posterior part, thereby explaining naturally the existence of animals having a caudal heart.