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PubMed2004

Gene expression during early ascidian metamorphosis requires signalling by Hemps, an EGF-like protein.

Hemps, a novel epidermal growth factor (EGF)-like protein, is expressed during larval development and early metamorphosis in the ascidian Herdmania curvata and plays a direct role in triggering metamorphosis. In order to identify downstream genes in the Hemps pathway we used a gene expression profiling approach, in which we compared post-larvae undergoing normal metamorphosis with larval metamorphosis blocked with an anti-Hemps antibody. Molecular profiling revealed that there are dynamic changes in gene expression within the first 30 minutes of normal metamorphosis with a significant portion of the genome (approximately 49%) being activated or repressed. A more detailed analysis of the expression of 15 of these differentially expressed genes through embryogenesis, larval development and metamorphosis revealed that while there is a diversity of temporal expression patterns, a number of genes are transiently expressed during larval development and metamorphosis. These and other differentially expressed genes were localised to a range of specific cell and tissue types in Herdmania larvae and post-larvae. The expression of approximately 24% of the genes that were differentially expressed during early metamorphosis was affected in larvae treated with the anti-Hemps antibody. Knockdown of Hemps activity affected the expression of a range of genes within 30 minutes of induction, suggesting that the Hemps pathway directly regulates early response genes at metamorphosis. In most cases, it appears that the Hemps pathway contributes to the modulation of gene expression, rather than initial gene activation or repression. A total of 151 genes that displayed the greatest alterations in expression in response to anti-Hemps antibody were sequenced. These genes were implicated in a range of developmental and physiological roles, including innate immunity, signal transduction and in the regulation of gene transcription. These results suggest that there is significant gene activity during the very early stages of H. curvata metamorphosis and that the Hemps pathway plays a key role in regulating the expression of many of these genes.

Animals↗

PubMed1999

Hemps, a novel EGF-like protein, plays a central role in ascidian metamorphosis.

All chordates share several characteristic features including a dorsal hollow neural tube, a notochord, a pharynx and an endostyle. Unlike other chordate taxa, ascidians have a biphasic life-history with two distinct body plans. During metamorphosis, the larval nerve cord and notochord degenerate and the pharyngeal gill slits and endostyle form. While ascidians, like other marine invertebrates, metamorphose in response to specific environmental cues, it remains unclear how these cues trigger metamorphosis. We have identified a novel gene (Hemps) which encodes a protein with a putative secretion signal sequence and four epidermal growth factor (EGF)-like repeats which is a key regulator of metamorphosis in the ascidian Herdmania curvata. Expression of Hemps increases markedly when the swimming tadpole larva becomes competent to undergo metamorphosis and then during the first 24 hours of metamorphosis. The Hemps protein is localised to the larval papillae and anterior epidermis of the larva in the region known to be required for metamorphosis. When the larva contacts an inductive cue the protein is released, spreading posteriorly and into the tunic as metamorphosis progresses. Metamorphosis is blocked by incubating larvae in anti-Hemps antibodies prior to the addition of the cue. Addition of recombinant Hemps protein to competent larvae induces metamorphosis in a concentration-dependent manner. A subgroup of genes are specifically induced during this process. These results demonstrate that the Hemps protein is a key regulator of ascidian metamorphosis and is distinct from previously described inducers of this process in terrestrial arthropods and aquatic vertebrates.

Animals↗

PubMed2001

Regulation of metamorphosis in ascidians involves NO/cGMP signaling and HSP90.

Treatment of larvae of the ascidians Boltenia villosa (Family: Pyuridae) and Cnemidocarpa finmarkiensis (Family: Styelidae) with drugs that inhibit the function of the molecular chaperone HSP90 increased the frequency of tail resorption, the primary morphogenetic event of metamorphosis. If treatment was initiated at hatching, metamorphic events subsequent to tail resorption failed to occur, indicating an ongoing role for HSP90 during morphogenesis. Removal of tails from heads of mature, but not newly hatched larvae, induced metamorphosis of the head. Decapitation experiments indicate that the capacity of tails to shorten in response to inhibition of HSP90 function requires communication with heads. To identify candidate proteins with which HSP90 may interact to regulate metamorphosis, we noted that in mammalian cells, nitric oxide synthase (NOS) interacts with HSP90 and its activity is sensitive to drugs that inhibit HSP90 function. In addition, nitric oxide (NO) signaling in the marine snail Ilyanassa obsoleta is an important regulator of metamorphosis. Inhibition of NOS activity in these ascidian larvae with L-NAME increased the frequency of metamorphosis, consistent with a putative interaction of NOS and HSP90. NOS is present in tail muscle cells, implicating them as targets for the drug treatments, consistent with the decapitation experiments. Inhibition of soluble guanylyl cyclase, the most common effector of NO signaling, also increased the frequency of metamorphosis. In contrast to treatment with anti-HSP90 drugs, metamorphosis induced with L-NAME or ODQ was complete. The results presented suggest that an HSP90-dependent, NO-based regulatory mechanism localized in tails represses ascidian metamorphosis. We discuss these results in relation to the induction of ascidian metamorphosis by several unrelated agents.

Animals↗

PubMed1986

Hormonal control of the intestinal brush border enzyme activities in developing anuran amphibians. I. Effects of hydrocortisone and insulin during and after spontaneous metamorphosis.

The effects of hydrocortisone and insulin on the intestinal brush border membrane enzymatic activities in an anuran amphibian, Alytes obstetricans, were investigated at the end of spontaneous metamorphosis and 2 weeks after its completion. At the end of metamorphosis, the brush border is differentiating in the apical region of a developing neoformed epithelium. Two weeks after the completion of metamorphosis, this epithelium is entirely formed. The animals received one hormone injection per day for 2 or 3 days running (hydrocortisone: 1, 5, or 25 micrograms/g body wt/day; insulin: 0.5, 1, or 5 mU/g body wt/day). The hydrolases studied were three glucosidases (maltase, glucoamylase, trehalase), gamma-glutamyl-transferase and alkaline phosphatase. In animals reaching the end of metamorphosis, hormonal treatments rarely modify the three glucosidase activities. Two weeks after metamorphosis, a 5 microgram/g body wt/day hydrocortisone injection usually results in a significant increase of the three glucosidase activities. Conversely, a 0.5 mU/g body wt/day insulin injection induced a marked decrease in these activities. At the end of metamorphosis, hydrocortisone has variable effects on gamma-glutamyl-transferase activity; insulin, however, does not significantly modify this activity. Two weeks later, insulin and sometimes hydrocortisone inhibit gamma-glutamyl-transferase activity. Whatever the developmental stage is, hydrocortisone is able to stimulate alkaline phosphatase activity. At the end of metamorphosis, insulin has no influence on this activity, but 2 weeks after metamorphosis, low doses of the hormone (0.5 mU/g body wt/day) significantly reduce it. These results emphasize the possibility that after spontaneous metamorphosis the enzymatic activities of the new intestinal brush border are hormone controlled. This control could be related to the development of the interrenal and pancreatic islet functions.

Alkaline Phosphatase↗

PubMed2004

Developmental expression and hormonal regulation of glucocorticoid and thyroid hormone receptors during metamorphosis in Xenopus laevis.

Corticosteroids, the primary circulating vertebrate stress hormones, are known to potentiate the actions of thyroid hormone in amphibian metamorphosis. Environmental modulation of the production of stress hormones may be one way that tadpoles respond to variation in their larval habitat, and thus control the timing of metamorphosis. Thyroid hormone and corticosteroids act through structurally similar nuclear receptors, and interactions at the transcriptional level could lead to regulation of common pathways controlling metamorphosis. To better understand the roles of corticosteroids in amphibian metamorphosis we analyzed the developmental and hormone-dependent expression of glucocorticoid receptor (GR) mRNA in the brain (diencephalon), intestine and tail of Xenopus laevis tadpoles. We compared the expression patterns of GR with expression of thyroid hormone receptor beta (TRbeta). In an effort to determine the relationship between nuclear hormone receptor expression and levels of ligand, we also analyzed changes in whole-body content of 3,5,3'-triiodothyronine (T(3)), thyroxine, and corticosterone (CORT). GR transcripts of 8, 4 and 2 kb were detected in all tadpole tissues, but only the 4 and 2 kb transcripts could be detected in embryos. The level of GR mRNA was low during premetamorphosis in the brain but increased significantly during prometamorphosis, remained at a constant level throughout metamorphosis, and increased to its highest level in the juvenile frog. GR mRNA level in the intestine remained relatively constant, but increased in the tail throughout metamorphosis, reaching a maximum at metamorphic climax. The level of GR mRNA was increased by treatment with CORT in the intestine but not in the brain or tail. TRbeta mRNA level increased in the brain, intestine and tail during metamorphosis and was induced by treatment with T(3). Analysis of possible crossregulatory relationships between GRs and TRs showed that GR mRNA was upregulated by exogenous T(3) (50 nM) in the tail but downregulated in the brain of premetamorphic tadpoles. Exogenous CORT (100 nM) upregulated TRbeta mRNA in the intestine. Our findings provide evidence for tissue-specific positive, negative and crossregulation of nuclear hormone receptors during metamorphosis of X. laevis. The synergy of CORT with T(3) on tadpole tail resorption may depend on the accelerated accumulation of GR transcripts in this tissue during metamorphosis, which may be driven by rising plasma thyroid hormone titers.

Animals↗

PubMed2001

Variable effects of goitrogens in inducing precocious metamorphosis in sea lampreys (Petromyzon marinus).

The ability of different goitrogens (anti-thyroid agents) to induce precocious metamorphosis in larval sea lampreys (Petromyzon marinus) was assessed in four separate experiments. Two of these goitrogens (propylthiouracil [PTU] and methimazole [MMI]) are inhibitors of thyroid peroxidase-catalyzed iodination, and three (potassium perchlorate [KClO(4)], potassium thiocyanate [KSCN], and sodium perchlorate [NaClO(4)]) are anionic competitors of iodide uptake. Because, theoretically, all of these goitrogens prevent thyroid hormone (TH) synthesis, we also measured their influence on serum concentrations of thyroxine and triiodothyronine. All goitrogens except PTU significantly lowered serum TH concentrations and induced metamorphosis in some larvae. The incidence of metamorphosis appeared to be correlated with these lowered TH concentrations in that KClO(4), NaClO(4), and MMI treatments resulted in the lowest serum TH concentrations and the highest incidence of metamorphosis in sea lampreys. Moreover, fewer larvae metamorphosed in the KSCN and low-KClO(4) treatment groups and their serum TH concentrations tended to be greater than the values in the aforementioned groups. MMI treatment at the concentrations used (0.087 and 0.87 mM) was toxic to 55% of the exposed sea lampreys within 6 weeks. The potassium ion administered as KCl did not alter serum TH concentrations or induce metamorphosis. On the basis of the results of these experiments, we have made the following conclusions: (i) In general, most goitrogens other than PTU can induce metamorphosis in larval sea lampreys, and this induction is coincident with a decline in serum TH concentrations. (ii) The method by which a goitrogen prevents TH synthesis is not directly relevant to the induction of metamorphosis. (iii) PTU has variable effects on TH synthesis and metamorphosis among lamprey species. (iv) Unlike in protochordates, potassium ions do not induce metamorphosis in sea lampreys and are not a factor in the stimulation of this event.

Animals↗

PubMed1995

Metamorphosis of spinal-projecting neurons in the brain of the sea lamprey during transformation of the larva to adult: normal anatomy and response to axotomy.

The spinal projecting system of the sea lamprey (Petromyzon marinus) has been used extensively in studies of axonal regeneration in both larvae and adults. However, little is known about the changes that are undergone by this system during metamorphosis. In order to determine the developmental changes in the size of the descending spinal projection and in the morphology of its neurons, larval, transforming, and adult lamprey brains were labeled by retrograde transport of horseradish peroxidase (HRP) injected into the spinal cord at 25% of body length. Examination of brain wholemount preparations revealed that the total number of labeled neurons doubled during metamorphosis. Most of this increase could be explained by elongation of reticulospinal axons from the rostralmost segments of the spinal cord to locations caudal to the injection site. There were no additions or deletions of either identified reticulospinal neurons or of reticulospinal nuclear groups between the larval and the adult stages. The proportions of Müller and Mauthner cells that were labeled reached a maximum of 93% during the early stages of metamorphosis. Axons of these neurons are known to project almost the entire length of the cord, even in larvae. Therefore, the efficiency of retrograde transport appears to be greater during metamorphosis than during larval or adult stages. While changes in efficiency of retrograde transport could account for some of the apparent increase in reticulospinal neuron numbers between larvae and animals undergoing metamorphosis, this could not contribute to the further increase in the apparent size of the reticulospinal system in the adult, since efficiency of retrograde labeling in these animals was lower than that at earlier stages. With retrograde labeling, a significant increase was seen in the profusion of dendritic arborization of some Müller and Mauthner cells during the early stages of metamorphosis. This correlated with an increase in the incidence of extreme axonal die-back, as indicated by the presence of retraction bulbs within the brainstem. However, intracellular injection of Neurobiotin in untransected animals showed similar degrees of dendritic arborization at all examined stages of development. Therefore, the dendritic profusion did not reflect developmental changes in neuronal morphology but rather reflected an increased sensitivity to axotomy during metamorphosis. We conclude that, during the transformation of the lamprey from the large larval to the adult form, there is little change in either the size or the dendritic morphology of the identified giant reticulospinal neurons. With respect to the smaller reticulospinal neurons, the distance of projection of many of their axons increases during metamorphosis, but there is very little increase in the number of reticulospinal neurons.

Animals↗

PubMed2002

Acceleration of Ambystoma tigrinum metamorphosis by corticotropin-releasing hormone.

Previous work of others and ours has shown that corticotropin-releasing hormone (CRH) is a positive stimulus for thyroid and interrenal hormone secretion in amphibian larvae and that activation of CRH neurons may mediate environmental effects on the timing of metamorphosis. These studies have investigated CRH actions in anurans (frogs and toads), whereas there is currently no information regarding the actions of CRH on metamorphosis of urodeles (salamanders and newts). We tested the hypothesis that CRH can accelerate metamorphosis of tiger salamander (Ambystoma tigrinum) larvae. We injected tiger salamander larvae with ovine CRH (oCRH; 1 microg/day; i.p.) and monitored effects on metamorphosis by measuring the rate of gill resorption. oCRH-injected larvae completed metamorphosis earlier than saline-injected larvae. There was no significant difference between uninjected and saline-injected larvae. Mean time to reach 50% reduction in initial gill length was 6.9 days for oCRH-injected animals, 11.9 days for saline-injected animals, and 14.1 days for uninjected controls. At the conclusion of the experiment (day 15), all oCRH-injected animals had completed metamorphosis, whereas by day 15, only 50% of saline-injected animals and 33% of uninjected animals had metamorphosed. Our results show that exogenous oCRH can accelerate metamorphosis in urodele larvae as it does in anurans. These findings suggest that the neuroendocrine mechanisms controlling metamorphosis are evolutionarily conserved across amphibian taxa.

Ambystoma↗

PubMed2003

Zebrafish puma mutant decouples pigment pattern and somatic metamorphosis.

The genetic and developmental bases for trait expression and variation in adults are largely unknown. One system in which genes and cell behaviors underlying adult traits can be elucidated is the larval-to-adult transformation of zebrafish, Danio rerio. Metamorphosis in this and many other teleost fishes resembles amphibian metamorphosis, as a variety of larval traits (e.g., fins, skin, digestive tract, sensory systems) are remodeled in a coordinated manner to generate the adult form. Among these traits is the pigment pattern, which comprises several neural crest-derived pigment cell classes, including black melanophores, yellow xanthophores, and iridescent iridophores. D. rerio embryos and early larvae exhibit a relatively simple pattern of melanophore stripes, but this pattern is transformed during metamorphosis into the more complex pattern of the adult, consisting of alternating dark (melanophore, iridophore) and light (xanthophore, iridophore) horizontal stripes. While it is clear that some pigment cells differentiate de novo during pigment pattern metamorphosis, the extent to which larval and adult pigment patterns are developmentally independent has not been known. In this study, we show that a subset of embryonic/early larval melanophores persists into adult stages in wild-type fish; thus, larval and adult pigment patterns are not completely independent in this species. We also analyze puma mutant zebrafish, derived from a forward genetic screen to isolate mutations affecting postembryonic development. In puma mutants, a wild-type embryonic/early larval pigment pattern forms, but supernumerary early larval melanophores persist in ectopic locations through juvenile and adult stages. We then show that, although puma mutants undergo a somatic metamorphosis at the same time as wild-type fish, metamorphic melanophores that normally appear during these stages are absent. The puma mutation thus decouples metamorphosis of the pigment pattern from the metamorphosis of many other traits. Nevertheless, puma mutants ultimately recover large numbers of melanophores and exhibit extensive pattern regulation during juvenile development, when the wild-type pigment pattern already would be completed. Finally, we demonstrate that the puma mutant is both temperature-sensitive and growth-sensitive: extremely severe pigment pattern defects result at a high temperature, a high growth rate, or both; whereas a wild-type pigment pattern can be rescued at a low temperature and a low growth rate. Taken together, these results provide new insights into zebrafish pigment pattern metamorphosis and the capacity for pattern regulation when normal patterning mechanisms go awry.

Animals↗

PubMed2005

Larval development and post-settlement metamorphosis of the barnacle Balanus albicostatus Pilsbry and the serpulid polychaete Pomatoleios kraussii Baird: Impact of a commonly used antifouling biocide, Irgarol 1051.

AbstractThe impact of a commonly-used antifouling algicide, Irgarol 1051, on the larval development and post-settlement metamorphosis of the barnacle, Balanus albicostatus Pilsbry (Crustacea: Cirripedia), and the larval metamorphosis of a serpulid polycheate, Pomatoleios kraussii Baird, was evaluated. In the case of B. albicostatus, larval mortality increased with an increase in the concentration of Irgarol 1051, and there was a shift in the larval stage targeted from advanced instars to early instars. Nauplii that survived to the cyprid instar stage when reared in the presence of Irgarol 1051 showed prolonged instar and total naupliar duration when compared to the controls. The post-settlement metamorphosis of cyprids significantly varied with Irgarol concentration and also with biofilm age. One and 2-d-old untreated biofilms showed higher metamorphosis when compared to 5-d-old biofilms. However, when the biofilms that promoted cyprid metamorphosis were treated with Irgarol 1051 at low concentrations, metamorphosis rates decreased. Cyprids were prevented from metamorphosing completely by biofilms treated at the highest concentration of Irgarol 1051. Inhibition of metamorphosis was also observed in the case of competent polychaete larvae when exposed to Irgarol 1051 compared to those exposed to metamorphosis inducers such as 3-iso-butyl-1-methylxanthine (IBMX) and natural biofilms. Identification of the pathway(s) that caused the promotory biofilms to become toxic when exposed to Irgarol 1051 is discussed.

1-Methyl-3-isobutylxanthine↗

PubMed2002

Timing is everything: the effects of putative dopamine antagonists on metamorphosis vary with larval age and experimental duration in the prosobranch gastropod Crepidula fornicata.

The signal transduction pathway through which excess potassium ion stimulates the larvae of many marine invertebrates to metamorphose is incompletely understood. Recent evidence suggests that dopamine plays important roles in the metamorphic pathway of Crepidula fornicata. Therefore, we asked whether blocking dopamine receptors might prevent excess potassium ion from stimulating metamorphosis in this species. Surprisingly, the effects of the three putative dopamine antagonists tested (all at 10 microM) varied with exposure duration and the age of competent larvae. Chlorpromazine, a nonspecific dopamine antagonist known to have a number of other pharmacological effects, blocked the inductive action of excess potassium ion during the initial 5-8-h exposure periods in most assays, particularly for younger or smaller competent larvae. However, chlorpromazine in the absence of excess potassium ion also stimulated metamorphosis, particularly over the next 18 h, and worked faster on older competent larvae than on younger competent larvae. The specific D(1) antagonist R(+)-Sch-23309 had similar effects, blocking potassium-stimulated metamorphosis in short-term exposures and stimulating metamorphosis in longer exposures, particularly for older competent larvae. Although the specific D(2) antagonist spiperone (SPIP) blocked the inductive effects of excess potassium ion in only 1 of 6 assays during the first 6 h of exposure, it blocked metamorphosis in 2 of the assays during 24-h exposures. Our results indicate that dopamine receptors are involved in the pathway through which excess potassium ion stimulates metamorphosis in C. fornicata. In addition, the largely latent inductive effects of chlorpromazine, an inhibitor of nitric oxide synthase, suggest that endogenous nitric oxide may play a natural role in inhibiting metamorphosis in this species. Overall, our results would then suggest that exposing larvae of C. fornicata to excess K(+) leads to a shutdown of nitric oxide synthesis via a dopaminergic pathway, a pathway that can be blocked by some dopamine antagonists. Alternatively, chlorpromazine might eventually be stimulating metamorphosis by elevating endogenous cyclic nucleotide (e.g., cAMP) concentrations, again acting downstream from the steps acted on directly by excess K(+).

Animals↗

PubMed1993

Calcium control of metamorphosis in polychaete larvae.

The importance of Ca2+ in the control of metamorphosis of a marine invertebrate larva was investigated. An excess of [Ca2+] in the external medium induced metamorphosis of Phragmatopoma californica (polychaete) larvae in a concentration-dependent manner. This effect is specific for calcium, and not simply the result of osmotic changes, as an excess of Mg2+ did not induce metamorphosis. Consistent with this finding, the calcium ionophore, A23187, also induced metamorphosis in a concentration-dependent manner. Paradoxically, however, the aromatic compounds diltiazem, verapamil, D600, and nifedipine, known to block Ca2+ channels in other systems, also induced metamorphosis. When exposed to diltiazem for only 20 h and subsequently washed free of this compound, 95% of the larvae metamorphosed and developed normally. Previous studies have demonstrated that the induction of metamorphosis in Phragmatopoma californica is controlled by chemosensory recognition of an exogenous morphogen and mediated by an excitatory pathway that involves adenyl cyclase and cyclic AMP. Because cellular excitation and cyclic AMP-dependent signal transduction generally involve the participation of calcium ion, the most parsimonious explanation for the results reported here include (1) direct control of the morphogenetic pathway by calcium ion, and (2) complexities of the calcium regulation of this process, or a functional similarity between the structurally related aromatic effectors tested and the natural inducer of metamorphosis.

Animals↗

PubMed2004

Ontogenic and ecological control of metamorphosis onset in a carapid fish, Carapus homei: experimental evidence from vertebra and otolith comparisons.

In Carapus homei, reef colonisation is associated with a penetration inside a sea cucumber followed by heavy transformations during which the length of the fish is reduced by 60%. By comparing vertebral axis to otolith ontogenetic changes, this study aimed (i) to specify the events linked to metamorphosis, and (ii) to establish to what extent these fish have the ability to delay it. Different larvae of C. homei were caught when settling on the reef and kept in different experimental conditions for at least 7 days and up to 21 days: darkness or natural light conditions, presence of sea cucumber or not, and food deprivation or not. Whatever the nutritional condition, a period of darkness seems sufficient to initiate metamorphosis. Twenty-one days in natural light conditions delayed metamorphosis, whereas the whole metamorphosis process is the fastest (15 days) for larvae living in sea cucumbers. Whether the metamorphosis was initiated or not, otoliths were modified with the formation of a transition zone, whose structure varied depending on the experimental conditions. At day 21, larvae maintained in darkness had an otolith transition zone with more increments (around 80), albeit wider than those (more or less 21) of individuals kept under natural lighting. These differences in otolith growth could indicate an increased incorporation rate of released metabolites by metamorphosing larvae. However, the presence of a transition zone in delayed-metamorphosis larvae suggests that these otolith changes record the endogenously-induced onset of metamorphosis, whereas body transformations seem to be modulated by the environmental conditions of settlement.

Animals↗

PubMed2005

Thyroid and pituitary gland development from hatching through metamorphosis of a teleost flatfish, the Atlantic halibut.

Fish larval development, not least the spectacular process of flatfish metamorphosis, appears to be under complex endocrine control, many aspects of which are still not fully elucidated. In order to obtain data on the functional development of two major endocrine glands, the pituitary and the thyroid, during flatfish metamorphosis, histology, immunohistochemistry and in situ hybridization techniques were applied on larvae of the Atlantic halibut (Hippoglossus hippoglossus), a large, marine flatfish species, from hatching through metamorphosis. The material was obtained from a commercial hatchery. Larval age is defined as day-degrees (D degrees =accumulated daily temperature from hatching). Sporadic thyroid follicles are first detected in larvae at 142 D degrees (27 days post-hatch), prior to the completion of yolk sack absorption. Both the number and activity of the follicles increase markedly after yolk sack absorption and continue to do so during subsequent development. The larval triiodothyronine (T(3)) and thyroxine (T(4)) content increases, subsequent to yolk absorption, and coincides with the proliferation of thyroid follicles. A second increase of both T(3) and T(4) occurs around the start of metamorphosis and the T(3) content further increases at the metamorphic climax. Overall, the T(3) content is lower than T(4). The pituitary gland can first be distinguished as a separate organ at the yolk sack stage. During subsequent development, the gland becomes more elongated and differentiates into neurohypophysis (NH), pars distalis (PD) and pars intermedia (PI). The first sporadic endocrine pituitary cells are observed at the yolk sack stage, somatotrophs (growth hormone producing cells) and somatolactotrophs (somatolactin producing cells) are first observed at 121 D degrees (23 days post-hatch), and lactotrophs (prolactin producing cells) at 134 D degrees (25 days post-hatch). Scarce thyrotrophs are evident after detection of the first thyroid follicles (142 D degrees ), but coincident with a phase in which follicle number and activity increase (260 D degrees ). The somatotrophs are clustered in the medium ventral region of the PD, lactotrophs in the anterior part of the PD and somatolactotrophs are scattered in the mid and posterior region of the pituitary. At around 600 D degrees , coinciding with the start of metamorphosis, somatolactotrophs are restricted to the interdigitating tissue of the NH. During larval development, the pituitary endocrine cells become more numerous. The present data on thyroid development support the notion that thyroid hormones may play a significant role in Atlantic halibut metamorphosis. The time of appearance and the subsequent proliferation of pituitary somatotrophs, lactotrophs, somatolactotrophs and thyrotrophs indicate at which stages of larval development and metamorphosis these endocrine cells may start to play active regulatory roles.

Animals↗

PubMed1987

Pharmacological evidence that alpha 1-adrenoceptors mediate metamorphosis of the Pacific oyster, Crassostrea gigas.

Oyster larvae can be induced to metamorphose by exposure to the natural vertebrate adrenergic agonists, epinephrine and norepinephrine. The larval receptors mediating this induction were pharmacologically characterized by testing the ability of a variety of adrenergic agonists and selected structural analogs of epinephrine and norepinephrine to induce oyster metamorphosis, and by testing the ability of various adrenergic antagonists to block the induction of metamorphosis by epinephrine. Oyster metamorphosis can be induced by vertebrate adrenergic agonists with relative potencies: cirazoline greater than epinephrine greater than phenylephrine greater than or equal to norepinephrine greater than alpha-methylnorepinephrine greater than isoproterenol much greater than methoxamine = clonidine. Other structural analogs of epinephrine and norepinephrine, including dopamine and octopamine, were ineffective at inducing metamorphosis. Induction of metamorphosis by epinephrine can be blocked by vertebrate adrenergic antagonists with relative potencies: chlorpromazine greater than or equal to prazosin greater than phentolamine greater than WB4101 greater than propranolol greater than yohimbine greater than metoprolol. These data demonstrate that receptors similar to vertebrate-type alpha 1-adrenoceptors mediate oyster metamorphosis. This is the first evidence for alpha 1-adrenoceptors in molluscs, and provides an important clue to the control of the complex process of molluscan metamorphosis and to the evolution of vertebrate adrenergic receptors.

Adrenergic alpha-Agonists↗

PubMed2005

Dopamine and serotonin modulate the onset of metamorphosis in the ascidian Phallusia mammillata.

Neurotransmitters play an important role in larval metamorphosis in different groups of marine invertebrates. In this work, the role of dopamine and serotonin during metamorphosis of the ascidian Phallusia mammillata larvae was examined. By immunofluorescence experiments, dopamine was localized in some neurons of the central nervous system and in the adhesive papillae of the larvae. Dopamine and serotonin signaling was inhibited by means of antagonists of these neurotransmitters receptors (R(+)-SCH-23390, a D(1) antagonist; clozapine, a D(4) antagonist; WAY-100635, a 5-HT(1A) antagonist) and by sequestering the neurotransmitters with specific antibodies. Moreover, dopamine synthesis was inhibited by exposing 2-cell embryos to alpha-methyl-l-tyrosine. Dopamine depletion, obtained by these different approaches, caused early metamorphosis, while serotonin depletion delayed the onset of metamorphosis. The opposite effects were obtained using agonists of the neurotransmitters: lisuride, a D(2) agonist, inhibited metamorphosis, while DOI hydrochloride and 8-OH-DPAT HBr, two serotonin agonists, promoted it. So, it is possible to suppose that dopamine signaling delayed metamorphosis while serotonin signaling triggers it. We propose a mechanism by which these neurotransmitters may modulate the timing of metamorphosis in larvae.

8-Hydroxy-2-(di-n-propylamino)tetralin↗

PubMed2001

Inhibition of coral fertilisation and larval metamorphosis by tributyltin and copper.

Fertilisation and larval metamorphosis of reef-building corals are important life history events leading to recruitment of juvenile corals to reef populations. Little is known of the sensitivity of these early life phases to pollution, or their relative susceptibility to certain toxicants compared with established coral colonies. Inhibition of fertilisation and larval metamorphosis of the coral Acropora millepora (Ehrenberg, 1834) was assessed in response to solutions of the antifoulants tributyltin (TBT) and copper (Cu) using laboratory-based bioassays. Nominal concentrations that inhibited 50% fertilisation and metamorphosis (IC50) were calculated from 4 h fertilisation and 24 h metamorphosis assays and were based on introduced dose. Cu was most potent towards fertilisation with an IC50 of 17.4 micrograms/l. TBT however, proved more toxic to larval metamorphosis having an IC50 of 2.0 micrograms/l. Inert surfaces coated with either Cu- or TBT-based antifouling paint also inhibited fertilisation and metamorphosis. The degree of inhibition was correlated with surface area of the paint coating. These results indicate fertilisation and metamorphosis of coral can be sensitive to active components of antifouling paints.

Animals↗

PubMed2001

Theory on the evolutionary history of lamprey metamorphosis: role of reproductive and thyroid axes.

Metamorphosis is a developmental strategy used by only a small number of extant fishes and little is known about its phylogenetic development during the evolution history of this large group of vertebrates. The present report provides a putative evolutionary history of metamorphosis in the lamprey, an extant agnathan with direct descendancy from some of the oldest known vertebrates. The study reviews recent data on the role of the thyroid gland and its hormones in metamorphosis, summarizes some recent views on the evolution of the endostyle/follicular thyroid in lampreys, and provides new data on the content of two gonadotropin-releasing hormones (GnRH-I and -III) in brain during goitrogen-stimulated, precocious metamorphosis. These new data support an earlier viewpoint of a relationship between thyroid and reproductive axes during metamorphosis. It is proposed that the earliest lampreys were paedomorphic larvae and they lived in a marine environment; as such, they resembled in many ways the larvae from which the ancient protochordates, Larvacea, are derived. The iodide-concentrating efficiency of the endostyle was a critical factor in the evolution of metamorphosis and this gland was replaced by a follicular thyroid, for postmetamorphic animals needed to store iodine following their invasion of freshwater. Larval growth and postmetamorphic reproduction in freshwater became fixtures in the lamprey life cycle; a non-parasitic adult life-history type appeared later. The presence among extant lampreys of two different adult life-history types, and examples of the lability of the timing of sexual maturation in some species, imply that there has been a complex interplay between the thyroid and reproductive axes during the evolution of metamorphosis in lampreys. This proposal is consistent with what we know of interplay of these axes in extant adult lampreys and with the long-held viewpoint that thyroid function and sexual maturation are an association with an ancient history.

Animals↗