The oldest gnathostome teeth.

Mandibular teeth and dentitions are features of jawed vertebrates that were first acquired by the Palaeozoic ancestors1-3 of living chondrichthyans and osteichthyans. The fossil record currently points to the latter part of the Silurian period4-7 (around 425 million years ago) as a minimum date for the appearance of gnathostome teeth and to the evolution of growth and replacement mechanisms of mandibular dentitions in the subsequent Devonian period2,8-10. Here we provide, to our knowledge, the earliest direct evidence for jawed vertebrates by describing Qianodus duplicis, a new genus and species of an early Silurian gnathostome based on isolated tooth whorls from Guizhou province, China. The whorls possess non-shedding teeth arranged in a pair of rows that demonstrate a number of features found in modern gnathostome groups. These include lingual addition of teeth in offset rows and maintenance of this patterning throughout whorl development. Our data extend the record of toothed gnathostomes by 14 million years from the late Silurian into the early Silurian (around 439 million years ago) and are important for documenting the initial diversification of vertebrates. Our analyses add to mounting fossil evidence that supports an earlier emergence of jawed vertebrates as part of the Great Ordovician Biodiversification Event (approximately 485-445 million years ago).

Neural origins of basal diencephalon in teleost fishes: Radial versus tangential migration.

Teleost fish possess large lateral diencephalic regions such as the torus lateralis, the preglomerular area, and the diffuse nucleus of the hypothalamic inferior lobe. While their developmental origins traditionally were suggested to lie in diencephalic midline ventricular proliferative zones, more remote midbrain origins were reported recently. This review focuses on the preglomerular region and summarizes the data supporting three existing hypotheses on its developmental origins. The conclusion is that lateral torus, diffuse nucleus of hypothalamic inferior lobe, and preglomerular region are part of the diencephalon, but have a multiregional origin provided by both radially and tangentially migrating cells forming these regions in question.

Bioinspired hierarchical impact tolerant materials.

The quest for new light-weight materials with superior mechanical properties is a goal of materials scientists and engineers worldwide. A promising route in this pursuit is drawing inspiration from nature to design and develop materials with enhanced properties. By emulating the graded mineral content and hierarchical structure of fish scales of the Arapaima gigas from the nano to macro scales, we were able to develop bioinspired laminated composites with improved impact resistance. Activated by the addition of nano-particles of Al2O3 and nano-layers of TiN to a thermoplastic fiber substrate, new energy dissipation mechanisms operating at the nanoscale enhanced the energy absorption and stiffness of the bioinspired material. Remarkably, the newly developed materials are easily transferred to the industry with minimum associated manufacturing costs.

Fluid-structure interaction modeling on a 3D ray-strengthened caudal fin.

In this paper, we present a numerical model capable of solving the fluid-structure interaction problems involved in the dynamics of skeleton-reinforced fish fins. In this model, the fluid dynamics is simulated by solving the Navier-Stokes equations using a finite-volume method based on an overset, multi-block structured grid system. The bony rays embedded in the fin are modeled as nonlinear Euler-Bernoulli beams. To demonstrate the capability of this model, we numerically investigate the effect of various ray stiffness distributions on the deformation and propulsion performance of a 3D caudal fin. Our numerical results show that with specific ray stiffness distributions, certain caudal fin deformation patterns observed in real fish (e.g. the cupping deformation) can be reproduced through passive structural deformations. Among the four different stiffness distributions (uniform, cupping, W-shape and heterocercal) considered here, we find that the cupping distribution requires the least power expenditure. The uniform distribution, on the other hand, performs the best in terms of thrust generation and efficiency. The uniform stiffness distribution, per se, also leads to 'cupping' deformation patterns with relatively smaller phase differences between various rays. The present model paves the way for future work on dynamics of skeleton-reinforced membranes.

Behavioral consequences of dietary exposure to crude oil extracts in the Siamese fighting fish (Betta splendens).

Uptake by fishes of crude oil and its polycyclic aromatic hydrocarbons (PAHs) components occurs via gills, dietary intake, or diffusion through the skin. Dietary exposure to crude oil and its components is environmentally relevant, and induces physiological and morphological disruptions in fish. However, the impacts of crude oil on fish social and reproductive behaviors and thus the possible influences on reproductive success are poorly understood. As a part of their intraspecific interactions, male Siamese fighting fish (Betta splendens) exhibit highly stereotypic behavioral and territorial displays. This makes this species a tractable model for testing crude oil effects on behavior. After 2 weeks of acclimation at 29 °C, male adult betta fish were divided into three groups and fed for 4 weeks with food spiked with water (control), low oil concentrations or high oil concentrations (∑Total PAH concentrations 340, 3960 or 8820 ng/g dw, respectively) to determine subsequent alterations in behavioral displays. Compared with control fish, the aggressive display of "opercular flaring" was significantly increased (P < 0.03, n = 14-16) in oil-exposed fish. Bubble nest building, as well as testis and brain mass, were significantly reduced in treated fish (P < 0.05). Hematocrit of treated groups was increased significantly (P < 0.02) from 21% in control fish to ∼27% in both oil exposure groups. Dietary exposure over a 4-week period to low, relevant levels of crude oil thus leads to an increase in aggressive behavioral displays, a decrease in reproductive activity and additional morphological changes.

Neurocranial anatomy of an enigmatic Early Devonian fish sheds light on early osteichthyan evolution.

The skull of 'Ligulalepis' from the Early Devonian of Australia (AM-F101607) has significantly expanded our knowledge of early osteichthyan anatomy, but its phylogenetic position has remained uncertain. We herein describe a second skull of 'Ligulalepis' and present micro-CT data on both specimens to reveal novel anatomical features, including cranial endocasts. Several features previously considered to link 'Ligulalepis' with actinopterygians are now considered generalized osteichthyan characters or of uncertain polarity. The presence of a lateral cranial canal is shown to be variable in its development between specimens. Other notable new features include the presence of a pineal foramen, the some detail of skull roof sutures, the shape of the nasal capsules, a placoderm-like hypophysial vein, and a chondrichthyan-like labyrinth system. New phylogenetic analyses place 'Ligulalepis' as a stem osteichthyan, specifically as the sister taxon to 'psarolepids' plus crown osteichthyans. The precise position of 'psarolepids' differs between parsimony and Bayesian analyses.

Imaging of Jurassic fossils from the Talbragar Fish Bed using fluorescence, photoluminescence, and elemental and mineralogical mapping.

The Talbragar Fish Bed is one of Australia's most important Jurassic deposits for freshwater fishes, land plants and aquatic and terrestrial insects. The site has yielded many well preserved fossils, which has led to the formal description of numerous new species and higher taxa. The excellent preservation of many fossils has allowed detailed anatomical studies, e.g. of the early teleost fish Cavenderichthys talbragarensis (Woodward, 1895). Here we report on the fluorescent characteristics and mineral composition of a range of Talbragar fossils. Most specimens fluoresce under ultraviolet, blue and green light. Elemental and mineralogical analyses revealed that the Talbragar fossils consist predominantly of quartz (SiO2), a mineral that is likely to account for the observed fluorescence, with trace kaolinite (Al2Si2O5(OH)4) in some of the fish fossils. Rock matrices are predominantly composed of quartz and goethite (FeO(OH)). Closer inspection of a plant leaf (Pentoxylon australicum White, 1981) establishes fluorescence as a useful tool for the visualisation of anatomical details that are difficult to see under normal light conditions.

Complex Systems Are More than the Sum of Their Parts: Using Integration to Understand Performance, Biomechanics, and Diversity.

Organisms are comprised of many interacting parts, and an increased number or specialization of those parts leads to greater complexity and the necessity for increased integration (the ability of those parts to perform together and maintain a functioning organism). Although this idea is widely recognized among biologists, organisms are more tangibly studied when those parts are considered independently. This reductionist approach has successfully advanced our understanding of organisms' performance. However, performance of one system might (or might not) be dependent on performance of another system to achieve a relevant outcome, and the mechanism of this dependence is poorly understood. We synthesize the concepts of complexity and integration and discuss their application in a biomechanical context. Capture of prey by predatory fishes is used as an example to highlight the application of these ideas. We provide a theoretical framework for future hypotheses of integration and predict an "integration space" for fishes that is then populated with data extracted from the literature. Additionally, using the kinematics of prey-capture in two species of sculpin (Scorpaeniformes: Cottidae), we show that species exhibit multivariate integration in distinct ways, and that these differences add additional insight into ecological divergence that would not be apparent by considering systems independently. Finally, we discuss new insights into organismal performance gained through the study of integration as an emergent property of kinematic systems working together during a common task. Integration is rarely the trait of interest, but we show that future work should adopt a more holistic approach to understand why and how animals perform complex behaviors.

Alterations of digestive enzyme activities, intestinal morphology and microbiota in juvenile paddlefish, Polyodon spathula, fed dietary probiotics.

The effects of dietary supplementation of probiotics on digestive enzymes activities, intestinal morphology and microbiota in juvenile paddlefish (Polyodon spathula) were studied. A total of 400 fish were reared in two cages and fed with a basal diet (control group, CG) or diet supplemented with commercial probiotics (treatment group, TG) for 80 days. Enzymes activities analysis indicated that protease and α-amylase activities increased (P < 0.01 or P < 0.05) in TG. Light microscopy observation demonstrated the decrease of wall thickness and muscularis thickness in foregut (P < 0.01), the increase of those in hindgut (P < 0.05), the increase of folds height in foregut (P < 0.01) and midgut in TG (P < 0.05). DGGE results of PCR-amplified 16S rRNA confirmed that the richness and diversity of intestinal microbial species increased in TG. The similarity between the commercial bacteria product and intestinal microbiota of TG were higher than the microbiota from CG. The quantities of bacterium, Firmicutes, Proteobacteria, Bacteroidetes, Fusobacteria, present an increasing trend from foregut to hindgut both in two groups. To our knowledge, this is the first in vivo study to reveal the effect of dietary probiotics on intestinal digestive enzymes activities, morphology and microbiota in paddlefish.

A suplementação de vitamina C na estruturação do tecido conjuntivo de melanotência-maçã / Vitamin C supplementation in the structuring of connective tissue of red rainbowfish

Peixes cultivados mostram-se altamente sensíveis a dietas deficientes em ácido ascórbico com sinais clínicos que afetam o desempenho e a comercialização por provocar deformidades. Com objetivo de conhecer os níveis mínimos para a melanotênia-maçã, foram testadas seis rações isoaminoacídicas e isocalóricas com níveis variados de vitamina C ativa. Ao final dos 42 dias experimentais, 36 peixes foram analisados histologicamente quanto à porcentagem de tecido conjuntivo e fibras musculares a fim de determinar a influência da porcentagem de Vitamina C nesses índices. Apesar de as médias serem estatisticamente iguais do ponto de vista quantitativo, qualitativamente os níveis mais altos de vitamina C possibilitaram melhor estrutura tecidual. A desestruturação do tecido muscular é um indicativo da deficiência alimentar com ácido ascórbico...

Farmed fishes are highly sensitive to diets deficient in ascorbic acid with clinical symptoms that affect performance and commercialization for causing deformities. Aiming to meet the minimum levels for red rainbow fish, six isoaminoacid and isocaloric diets with varying levels of active vitamin C were tested. At the end of 42 days experimental period, 36 fishes were histologically analyzed for the percentage of connective tissue and muscle fibers in order to determine the influence of the Vitamin C percentage in those ratios. Although the averages are statistically indistinguishable from the quantitative point of view, qualitatively higher levels of vitamin C allow better tissue structure. The breakdown of muscle tissue is indicative for dietary deficiency of ascorbic acid...

The vertebrate diencephalic MCH system: a versatile neuronal population in an evolving brain.

Neurons synthesizing melanin-concentrating hormone (MCH) are described in the posterior hypothalamus of all vertebrates investigated so far. However, their anatomy is very different according to species: they are small and periventricular in lampreys, cartilaginous fishes or anurans, large and neuroendocrine in bony fishes, or distributed over large regions of the lateral hypothalamus in many mammals. An analysis of their comparative anatomy alongside recent data about the development of the forebrain, suggests that although very different, MCH neurons of the caudal hypothalamus are homologous. We further hypothesize that their divergent anatomy is linked to divergence in the forebrain - in particular telencephalic evolution.

The earliest known stem-tetrapod from the Lower Devonian of China.

Recent discoveries of advanced fish-like stem-tetrapods (for example, Panderichthys and Tiktaalik) have greatly improved our knowledge of the fin-to-limb transition. However, a paucity of fossil data from primitive finned tetrapods prevents profound understanding of the acquisition sequence of tetrapod characters. Here we report a new stem-tetrapod (Tungsenia paradoxa gen. et sp. nov.) from the Lower Devonian (Pragian, ∼409 million years ago) of China, which extends the earliest record of tetrapods by some 10 million years. Sharing many primitive features with stem-lungfishes, the new taxon further fills in the morphological gap between tetrapods and lungfishes. The X-ray tomography study of the skull depicts the plesiomorphic condition of the brain in the tetrapods. The enlargement of the cerebral hemispheres and the possible presence of the pars tuberalis in this stem-tetrapod indicate that some important brain modifications related to terrestrial life had occurred at the beginning of the tetrapod evolution, much earlier than previously thought.

The body weight changes and gut morphometry of clarias gariepinus juveniles on feeds supplemented with walnut (Tetracarpidium conophorum) leaf and onion (Allium cepa) bulb residues / Cambios de peso corporal y morfometría intestinal en clarias gariepinus juvenilespor alimentos suplementados con hojas de nogal (Tetracarpidium conophorum)y residuos de bulbo de cebolla (Allium cepa)

This experiment was conducted to evaluate the effects of walnut (T. conophorum) leaf and onion (A. cepa) bulb on the gut morphometry of Clarias gariepinus towards elucidating its mechanism of growth promotion. Twenty Clarias gariepinus juveniles (7.39+/-0.29g/ fish) were allocated to nine treatments, with three replicates each. Nine experimental diets were formulated at 40 percent crude protein representing different level of walnut leaf and onion bulb (0.5 percent, 1.0 percent, 1.5 percent and 2.0 percent) respectively and the control diet was free from walnut leaf and onion bulb and were fed twice daily at 3 percent body weight for 12 weeks. Growth indices like Mean Weight Gain (MWG), Specific Growth Rate (SGR), Feed Intake (FI) and Feed Conversion Ratio (FCR) were measured. Gut morphormetry such as villi length, villi width and cryptal depth were investigated. Data were analyzed using linear regression, descriptive statistics and ANOVA at p= 0.05. Results showed that weight gain and specific growth rate of Clarias gariepinus increased with increasing level of walnut leaf and onion bulb residues. The highest weight gain (53.81+/-5.85), specific growth rate (1.09+/-0.11), feed conversion ratio (2.16+/-0.02) was observed with 1.5 percent inclusions of walnut leaf residue diets. Although, FCR and SGR were not significantly different (p>0.05) among the treatments, the treated groups had better area of absorption and cryptal depth compared to the control. Treatment with 1.5 percent inclusion of walnut leaf residue recorded highest villi length and villi width and treatment with 1.5 percent inclusion of onion bulb recorded highest cryptal depth (30.00+/-0.00). These results indicate that using walnut leaf and onion bulb as a supplement in plant ­based diets may be useful in improving feed acceptability and growth performance of cultured C. gariepinus fed low cost, plant ­ based diets and it was concluded that walnut (T. conophorum) leaf at 1.5 percent inclusion...

Este estudio fue realizado para evaluar los efectos de la hoja del nogal (T. conophorum) y el bulbo de la cebolla (A. cepa) sobre la morfometría intestinal de Clarias gariepinus con el fin de dilucidar su mecanismo de promoción del crecimiento. Veinte Clarias gariepinus jóvenes (7,39+/-0,29 g/pez) fueron asignadas a nueve tratamientos, con tres repeticiones cada uno. Nueve dietas experimentales se formularon con la proteína cruda al 40 por ciento que representan los diferentes niveles de hoja del nogal y el bulbo de la cebolla (0,5 por ciento, 1,0 por ciento, 1,5 por ciento y 2,0 por ciento), respectivamente, y dieta de control libre de la hoja del nogal y el bulbo de la cebolla, que fueron alimentados dos veces al día en un 3 por ciento del peso corporal durante 12 semanas. Fueron medidos los índices de crecimiento como aumento de peso medio (APM), tasa de crecimiento específico (TCE), consumo de alimento (CA) y el índice de conversión alimeticia (ICA). La morfometría intestinal como longitud de las vellosidades, ancho de las vellosidades y profundidad de las criptas fueron investigados. Los datos fueron analizados mediante regresión lineal, estadística descriptiva y ANOVA con un valos p=0,05. Los resultados mostraron que el aumento de peso y la tasa de crecimiento específico de Clarias gariepinus se elevó al aumentar el nivel de la hoja de nogal y los residuos del bulbo de cebolla. La mayor ganancia de peso (53,81 +/- 5,85), tasa de crecimiento específico (1,09 +/- 0,11), tasa de conversión alimenticia (2,16 +/- 0,02) se observó con inclusiones de 1,5 por ciento de las dietas de residuos de hojas de nogal. Aunque el ICA y TCE no fueron significativamente diferentes (p> 0,05) entre los tratamientos. Los grupos tratados tuvieron una mejor área de absorción y profundidad de las criptas en comparación con el control. El tratamiento con 1,5 por ciento de inclusión de hojas de nogal registró la mayor longitud y ancho de las vellosidades y el tratamiento con 1,5...

Organization of the cholinergic systems in the brain of two lungfishes, Protopterus dolloi and Neoceratodus forsteri.

Lungfishes (dipnoans) are currently considered the closest living relatives of tetrapods. The organization of the cholinergic systems in the brain has been carefully analyzed in most vertebrate groups, and major shared characteristics have been described, although traits particular to each vertebrate class have also been found. In the present study, we provide the first detailed information on the distribution of cholinergic cell bodies and fibers in the central nervous system in two representative species of lungfishes, the African lungfish (Protopterus dolloi) and the Australian lungfish (Neoceratodus forsteri), as revealed by immunohistochemistry against the enzyme choline acetyltransferase (ChAT). Distinct groups of ChAT immunoreactive (ChAT-ir) cells were observed in the basal telencephalon, habenula, isthmic nucleus, laterodorsal tegmental nucleus, cranial nerve motor nuclei, and the motor column of the spinal cord, and these groups seem to be highly conserved among vertebrates. In lungfishes, the presence of a cholinergic cell group in the thalamus and the absence of ChAT-ir cells in the tectum are variable traits, unique to this group and appearing several times during evolution. Other characters were observed exclusively in Neoceratodus, such as the presence of cholinergic cells in the suprachiasmatic nucleus, the pretectal region and the superior raphe nucleus. Cholinergic fibers were found in the medial pallium, basal telencephalon, thalamus and prethalamus, optic tectum and interpeduncular nucleus. Comparison of these results with those from other classes of vertebrates, including a segmental analysis to correlate cell populations, reveals that the cholinergic systems in lungfishes largely resemble those of amphibians and other tetrapods.

Faking giants: the evolution of high prey clearance rates in jellyfishes.

Jellyfishes have functionally replaced several overexploited commercial stocks of planktivorous fishes. This is paradoxical, because they use a primitive prey capture mechanism requiring direct contact with the prey, whereas fishes use more efficient visual detection. We have compiled published data to show that, in spite of their primitive life-style, jellyfishes exhibit similar instantaneous prey clearance and respiration rates as their fish competitors and similar potential for growth and reproduction. To achieve this production, they have evolved large, water-laden bodies that increase prey contact rates. Although larger bodies are less efficient for swimming, optimization analysis reveals that large collectors are advantageous if they move through the water sufficiently slowly.

Ontogeny of swimming speed, schooling behaviour and jellyfish avoidance by Japanese anchovy Engraulis japonicus.

The ontogeny of swimming speed, schooling behaviour and jellyfish avoidance was studied in hatchery-reared Japanese anchovy Engraulis japonicus to compare its life-history strategy with two other common pelagic fishes, jack mackerel Trachurus japonicus and chub mackerel Scomber japonicus. Cruise swimming speed of E. japonicus increased allometrically from 1·4 to 3·9 standard length (L(S) ) per s (L(S) s(-1) ) from early larval to metamorphosing stage. Burst swimming speed also increased from 6·1 to 28 L(S) s(-1) in these stages. Cruise speed was inferior to that of S. japonicus, as was burst speed to that of T. japonicus. Engraulis japonicus larvae were highly vulnerable to predation by moon jellyfish Aurelia aurita and were readily eaten until they reached 23 mm L(S) , but not at 26 mm L(S) . Schooling behaviour (indicated by parallel swimming) started at c. 17 mm L(S) . Average distance to the nearest neighbour was shorter than values reported in other pelagic fishes. The relatively low predator avoidance capability of E. japonicus may be compensated for by their transparent and thus less conspicuous body, in addition to their early maturation and high fecundity.

Tetrapod trackways from the early Middle Devonian period of Poland.

The fossil record of the earliest tetrapods (vertebrates with limbs rather than paired fins) consists of body fossils and trackways. The earliest body fossils of tetrapods date to the Late Devonian period (late Frasnian stage) and are preceded by transitional elpistostegids such as Panderichthys and Tiktaalik that still have paired fins. Claims of tetrapod trackways predating these body fossils have remained controversial with regard to both age and the identity of the track makers. Here we present well-preserved and securely dated tetrapod tracks from Polish marine tidal flat sediments of early Middle Devonian (Eifelian stage) age that are approximately 18 million years older than the earliest tetrapod body fossils and 10 million years earlier than the oldest elpistostegids. They force a radical reassessment of the timing, ecology and environmental setting of the fish-tetrapod transition, as well as the completeness of the body fossil record.

Distribution of androgen receptor mRNA expression in vocal, auditory, and neuroendocrine circuits in a teleost fish.

Across all major vertebrate groups, androgen receptors (ARs) have been identified in neural circuits that shape reproductive-related behaviors, including vocalization. The vocal control network of teleost fishes presents an archetypal example of how a vertebrate nervous system produces social, context-dependent sounds. We cloned a partial cDNA of AR that was used to generate specific probes to localize AR expression throughout the central nervous system of the vocal plainfin midshipman fish (Porichthys notatus). In the forebrain, AR mRNA is abundant in proposed homologs of the mammalian striatum and amygdala, and in anterior and posterior parvocellular and magnocellular nuclei of the preoptic area, nucleus preglomerulosus, and posterior, ventral and anterior tuberal nuclei of the hypothalamus. Many of these nuclei are part of the known vocal and auditory circuitry in midshipman. The midbrain periaqueductal gray, an essential link between forebrain and hindbrain vocal circuitry, and the lateral line recipient nucleus medialis in the rostral hindbrain also express abundant AR mRNA. In the caudal hindbrain-spinal vocal circuit, high AR mRNA is found in the vocal prepacemaker nucleus and along the dorsal periphery of the vocal motor nucleus congruent with the known pattern of expression of aromatase-containing glial cells. Additionally, abundant AR mRNA expression is shown for the first time in the inner ear of a vertebrate. The distribution of AR mRNA strongly supports the role of androgens as modulators of behaviorally defined vocal, auditory, and neuroendocrine circuits in teleost fish and vertebrates in general.

Distribution of somatostatin immunoreactive neurons and fibres in the central nervous system of a chondrostean, the Siberian sturgeon (Acipenser baeri).

Somatostatin (SOM) is a neuropeptide that is widely distributed in the central nervous system of vertebrates. Two isoforms of somatostatin (SS1 and SS2) have been characterized in sturgeon and in situ hybridisation studies in the sturgeon brain have demonstrated that mRNAs of the two somatostatin precursors (PSS1 and PSS2) are differentially expressed in neurons [Trabucchi, M., Tostivint, H., Lihrmann, I., Sollars, C., Vallarino, M., Dores, R.M., Vaudry, H., 2002. Polygenic expression of somatostatin in the sturgeon Acipenser transmontanus: molecular cloning and distribution of the mRNAs encoding two somatostatin precursors. J. Comp. Neurol. 443, 332-345.]. However, neither the morphology of somatostatinergic neurons nor the patterns of innervation have yet been characterized. To gain further insight into the evolution of this system in primitive bony fishes, we studied the distribution of somatostatin-immunoreactive (SOM-ir) cells and fibres in the brain of the Siberian sturgeon (Acipenser baeri). Most SOM-ir cells were found in the preoptic area and hypothalamus and abundant SOM-ir fibres coursed along the hypothalamic floor towards the median eminence, suggesting a hypophysiotrophic role for SOM in sturgeon. In addition, SOM-ir cells and fibres were observed in extrahypothalamic regions such as the telencephalon thalamus, rhombencephalon and spinal cord, which also suggests neuromodulatory and/or neurotransmitter functions for this peptide. Overall there was a good correlation between the distribution of SOM-ir neurons throughout the brain of A. baeri and that of PSS1 mRNA in Acipenser transmontanus. Comparative analysis of the results with those obtained in other groups of fishes and tetrapods indicates that widespread distribution of this peptide in the brain is shared by early vertebrate lines and that the general organization of the somatostatinergic systems has been well-conserved during evolution.