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PubMed1978

The identification of an octopaminergic neurone and the modulation of a myogenic rhythm in the locust.

An octopaminergic neurone in an insect is demonstrated. This cell (DUMETi) is the dorsal unpaired median neurone which projects to the extensor tibiae muscle of the locust metathoracic leg. Its soma was physiologically identified, isolated and shown to contain about 0.1 pmol of octopamine. Octopamine is about four times more concentrated in the axon than in the soma. The concentration in the soma is at least 800 times more than that in the soma of an identified motoneurone (which controls the fast extensor of the tibia). The effects of DUMETi on a myogenic rhythm in the extensor muscle of the locust metathoracic leg can be mimicked by superfusion with low concentrations of octopamine. The myogenic bundle possesses at least two types of aminergic receptor: one which slows the rhythm (and has a high-affinity for octopamine) and a second which accelerates the rhythm (and has a low-affinity for octopamine but a high-affinity for the indolalkylamine, 5-hydroxytryptamine). The roles of the two receptor types in modulating the rhythm are discussed in relation to the function of the rhythm.

Action Potentials↗
PubMed2003

Modulation of early olfactory processing by an octopaminergic reinforcement pathway in the honeybee.

Processing of olfactory information in the antennal lobes of insects and olfactory bulbs of vertebrates is modulated by centrifugal inputs that represent reinforcing events. Octopamine release by one such pathway in the honeybee antennal lobe modulates olfactory processing in relation to nectar (sucrose) reinforcement. To test more specifically what role octopamine plays in the antennal lobe, we used two treatments to disrupt an octopamine receptor from Apis mellifera brain (AmOAR) function: (1) an OAR antagonist, mianserin, was used to block receptor function, and (2) AmOAR double-stranded RNA was used to silence receptor expression. Both treatments inhibited olfactory acquisition and recall, but they did not disrupt odor discrimination. These results suggest that octopamine mediates consolidation of a component of olfactory memory at this early processing stage in the antennal lobe. Furthermore, after consolidation, octopamine release becomes essential for recall, which suggests that the modulatory circuits become incorporated as essential components of neural representations that activate odor memory.

Amino Acid Sequence↗
PubMed1992

The octopaminergic system within the ventral nerve cord of the American cockroach.

Octopamine-immunoreactive neurons within the ventral nerve cord of the cockroach, Periplaneta americana, were mapped with a new anti-octopamine serum. The specificity of this antiserum was demonstrated by dot blot immunoassay and by comparing the immunocytochemical staining patterns obtained after incubation with anti-dopamine and anti-octopamine serum. Putative octopaminergic dorsal and ventral unpaired median (DUM resp. VUM) neurons showed octopamine-like immunoreactivity in all ventral ganglia. The numbers of DUM cells in the mesothoracic, metathoracic and terminal ganglia of females correspond to those previously characterized by retrograde staining 19, 33, 34. It could be shown that besides segmentally projecting there are also intersegmentally projecting DUM neurons within the thoracic ganglia. In addition various, previously unknown, paired octopamine-ir cells were revealed in all ventral ganglia except the abdominal ganglia 2-5.

Animals↗
PubMed1986

Role of ascorbic acid in dopamine beta-hydroxylation. The endogenous enzyme cofactor and putative electron donor for cofactor regeneration.

The role(s) of ascorbic acid in dopamine beta-hydroxylation was studied in primary cultures of bovine adrenomedullary chromaffin cells and in isolated bovine adrenomedullary chromaffin vesicles. Dopamine beta-hydroxylase activity was assessed by measuring the rate of conversion of tyramine to octopamine. The ascorbic acid content of chromaffin cells declined with time in culture and the dopamine beta-hydroxylase activity of ascorbate-depleted cells was low. Ascorbate additions to ascorbate-depleted cells increased both the intracellular ascorbate concentrations and the rates of dopamine beta-hydroxylation. Ascorbate uptake into the cells was rapid; however, the onset of enhanced octopamine synthesis by added ascorbate was delayed by several hours and closely followed the time course for accumulation of the newly taken up ascorbate into the chromaffin vesicle. The amount of octopamine synthesized by the chromaffin cells exceeded the intracellular ascorbate content and ascorbate levels were maintained during dopamine beta-hydroxylation in the absence of external ascorbate. This suggests an efficient recycling of ascorbate. In contrast to intact cells, ascorbic acid was depleted during octopamine synthesis in isolated chromaffin vesicles. The molar ratio of octopamine formed to ascorbate depleted was close to unity. Thus, the recycling of intravesicular ascorbate depends on an extravesicular factor(s). The depletion of intravesicular ascorbate during dopamine beta-hydroxylation was prevented by the addition of nonpermeant extravesicular electron donors such as ascorbate or glucoascorbate. This suggests that intravesicular ascorbate is maintained in the reduced state by electron transport across the vesicle membrane. These results are compatible with the hypothesis that both intra- and extravesicular ascorbate participate in the regulation of dopamine beta-hydroxylase. Intravesicular ascorbate is the cofactor for the enzyme. Cytosolic ascorbate is most likely the electron donor for the vesicle-membrane electron transport system which maintains the intravesicular cofactor concentration.

Adrenal Medulla↗
PubMed1982

The pharmacology of Limulus central neurons.

1. Intracellular recordings have been made from neurons in the central nervous system of the horse-shoe crab, Limulus polyphemus. Neurons possess resting potentials between -40 and -60 mV, with action potentials ranging from 2-3 mV up to 60 mV in amplitude. Neurons also have excitatory and inhibitory postsynaptic potentials. 2. All the neurons studied are inhibited by GABA and excited by cholinomimetics. The GABA response is chloride mediated and reversibly antagonised by picrotoxinin but not by bicuculline or bicuculline methochloride or methoiodide. The cholinergic response is nicotinic and blocked by pentolinium, hexamethonium, chlorisondamine and dihydro-beta-erythroidine. 3. L-Glutamate can excite some cells, inhibit others and have a biphasic action, inhibition followed by excitation, on other cells. The inhibitory effect is chloride mediated and blocked by picrotoxinin. Ibotenate mimics the action of glutamate both in terms of inhibition and excitation but kainate and quisqualate only mimic the excitatory action of L-glutamate. 4. Dopamine, octopamine, 5-hydroxytryptamine and histamine excite some neurons while inhibiting others or have a biphasic action. Dopamine and octopamine normally have different effects on the same cell, suggesting they act via different receptors. Octopamine shows stereospecificity for the (-) isomer which is more than 100 times more active than the (+) isomer and octopamine is reversibly antagonised by phentolamine and cyproheptadine. 5. Proctolin has an excitatory action on these neurons and this effect is long lasting and can be potentiated by dibutyl cyclic AMP. 6. The pharmacology of Limulus central neurons is compared to the pharmacology of insect and crustacean central neurons. It is concluded that GABA and acetylcholine are central transmitters throughout the arthropods. It is also probable that L-glutamate and octopamine have a physiological role in the arthropod central nervous system. Proctolin appears to modify neuronal and muscle activity in the arthropods and has a modulatory or transmitter function.

Acetylcholine↗
PubMed1993

The effect of the inactive mutation on longevity, sex, rhythm and resistance to p-cresol in Drosophila melanogaster.

Flies carrying the inactive (iav) mutation exhibit low locomotor activity and poor mating success, both of which are associated with a deficiency in the putative neurotransmitter, octopamine. Several other aspects of the iav mutant phenotype are described here. Male and female iav mutants show a small reduction in longevity but it is not clear whether this is a consequence of the iav mutation or their inactive phenotype. Young iav males show extended attractiveness to older courting males, which supports the notion that the iav gene has a role in post-eclosional maturation. The eclosion rhythm of iav mutants is normal, discounting the possibility of a role for octopamine in the maintenance of circadian rhythm. Flies carrying the iav mutation are highly susceptible to the octopamine analogue p-Cresol. Other phenotypically inactive flies show wild type levels of p-Cresol resistance. This is attributed to the deficiency of octopamine in iav mutants because low octopamine levels may be unable to out-compete the toxic effect of p-Cresol. Some inferences on the possible mode of action of the iav gene product are discussed.

Animals↗
PubMed1987

cAMP elevation modulates physiological activity of pyloric neurons in the lobster stomatogastric ganglion.

1. We analyzed the physiological effects of the adenylate cyclase activator forskolin, and other adenosine 3',5'-cyclic monophosphate (cAMP)-elevating agents, on neurons of the pyloric circuit from the stomatogastric ganglion of the lobster Panulirus interruptus. Agents were bath applied to pyloric neurons either in the synaptically intact pyloric circuit or following isolation from all known synaptic input. 2. Several cAMP-elevating agents, including forskolin, 3-isobutyl-1-methylxanthine, Ro20-1724, and 8-bromo-cAMP, generated similar motor patterns from the pyloric circuit. The motor patterns exhibited an increased cycle frequency and enhanced spike activity from all classes of pyloric neurons. Since these agents differ both in structure and site of action in the cAMP pathway, their physiological effects on the motor pattern probably result from increased cAMP levels in pyloric neurons. 3. When forskolin was applied to synaptically isolated neurons, it caused a strong activation or enhancement of activity of all pyloric cells. However, it induced different types of activity in different cells, including the induction of bursting pacemaker potentials in one cell type, activation of plateau potentials in another, and depolarization with activation or enhancement of tonic spike activity in the remaining cells. Thus there is no single physiological response to cAMP elevation in the pyloric circuit; its effects can be quite diverse, mediating several activity states, in different cells. 4. Radioimmunoassays were performed on whole stomatogastric ganglia to determine whether known neuromodulators can affect cAMP concentrations. Both forskolin and octopamine increased cAMP levels, whereas dopamine, serotonin, proctolin, and FMRFamide did not appreciably affect cAMP levels. The physiological effects of octopamine and forskolin are similar in most, but not all, pyloric cells. Octopamine is thus a candidate neuromodulator whose actions may be mediated, at least in part, by increased cAMP in some pyloric cells; however, forskolin does not completely mimic the physiological effects of octopamine on all pyloric neurons, suggesting that octopamine can also act by other biochemical mechanisms.

1-Methyl-3-isobutylxanthine↗
PubMed1997

Octopaminergic innervation and modulation of a locust flight steering muscle

We demonstrate that the meso- and metathoracic pleuroaxillary flight steering muscle (M85 mesothorax, M114 metathorax) of the migratory locust are each innervated by a single dorsal unpaired median neurone (DUM3,4,5a). The soma of this neurone can be localized by retrograde staining of the motor nerve with Neurobiotin, but not with cobalt salts. The primary neurite projects in the superficial DUM cell tract, and the axons run in nerve roots 3, 4 and 5 and in all their secondary branches. Other muscle targets include the second tergal remotor coxa (M120) and the posterior rotator coxae (M122, M123, M124), but not the first tergal remotor coxa (M119) and subalar (M129) flight muscles. Octopamine-like immunoreactive varicosities occur on the pleuroaxillary muscles. Stimulation of DUM3,4,5a and octopamine (10(-6) mol l-1) superfusion increased the amplitude and the relaxation velocity of neurally evoked twitch contractions of this muscle. Octopamine also significantly reduced the tonic tension that this muscle develops when stimulated at flight frequency (20 Hz), while increasing the amplitude of each phasic twitch. A catch-like tension is also reduced in the presence of octopamine. Simulations of the motor pattern experienced by the pleuroaxillary muscles during roll manoeuvres suggest that transient changes in tension underlying corrective steering could be doubled in the presence of octopamine.

Journal Article↗
PubMed1999

In vitro and in vivo effects of formamidines in locust (Locusta migratoria migratorioides).

In vivo and in vitro experiments were used to study the effects of formamidines in the locust, Locusta migratoria migratorioides. In vivo the lethal and the antifeeding effects, in vitro the inhibition of the binding of a selective 3H-ligand to the receptors of octopamine, tyramine, dopamine, serotonin and gamma-amino butiric acid were studied. We have demonstrated that demethylchlordimeform is specific agonist to octopamine receptor, having high affinity to octopamine receptor, a moderate affinitiy to tyramine receptor and a low affinity to dopamine, serotonin and to gamma-amino butiric acid receptors. The demethylated chlordimeform analogoues, demethylchlordimeform and didemethylchlordimeform have higher affinity to the octopamine receptor than the parent compound. The formamidines had a toxic and an antifeeding effects when injected into the locust. The half lethal doses (LD50) and the feeding inhibition were correlated with the affinity of the compounds (Ki). The ring substitutions of the mulecule have alterated the both affinity and in vivo effect of the compounds. The most effective ring substitution pattern is 2,4-disubstitution with a combination of methyl groups or halogens. Our results suggest that the lethal effect of formamidines is mediated through the octopamine receptor.

Amidines↗
PubMed1981

Pathogenesis of hepatic encephalopathy and hyperdynamic syndrome in cirrhosis. Role of false neurotransmitters.

We sought to determine whether false neurotransmitters (FNTs) play an important role as determinants not only of hepatic encephalopathy but also of hyperdynamic syndrome in cirrhosis. A combined biochemical and hemodynamics study of 55 bleeding cirrhotic patients was made. We evaluated the aromatic and aliphatic branched-chain amino acids and octopamine serum levels as well as the hemodynamic measurements. The results show that there is a correlation between levels of serum octopamine and aromatic amino acids and hepatic coma: the higher the octopamine level, the deeper the hepatic coma. There is also a correlation between aromatic amino acids and cardiac index and total peripheral resistance. Furthermore, when a narrowing of arteriovenous difference in oxygen occurs and oxygen consumption decreases, there is an increase not only in the level of aromatic amino acids, but also in octopamine level, suggesting an important linkage between hemodynamic and metabolic impairment.

Adolescent↗
PubMed1993

Neurotransmitters in the nervous system of Macoma balthica (Bivalvia).

The distribution of histamine-, octopamine-, gamma-aminobutyric acid- (GABA) and taurine-like immunoreactivity in the bivalve mollusc Macoma balthica was studied immunocytochemically with antisera produced in rabbits. Histamine levels in the ganglia and whole animals were also measured by high-performance liquid chromatography using a postcolumn derivatization method. Immunoreactivity for these substances, except for taurine, is found in the central nervous system of this species. The most extensive neuronal system is revealed with the antiserum against histamine. All the main ganglia contain histamine-immunoreactive cell bodies, and a dense network of nerve fibers is seen in the ganglia and nerve roots. Histamine-immunoreactive nerve fibers project to the mantle edge, lips and oesophagus. The basal part of the inhalant siphon is rich in histamine-immunoreactive fibers. Unlike histamine, octopamine- and GABA-like immunoreactivities are restricted to the central nervous system. Taurine-like immunoreactivity is not found in the nervous system of this species. In the nervous system, histamine-immunoreactive cell bodies and fibers are more numerous than those that are octopamine- and GABA-immunoreactive. The distribution of these substances in the ganglia is different. GABA-immunoreactive cells are typically smaller than most of the histamine- and octapamine-immunoreactive cells. Most GABA- and octopamine-immunoreactive cells and fibers are located in the pedal ganglion. Histamine is distributed more evenly in the ganglia and nerve roots. The biochemical measurements of histamine correlate well with the immunohistochemical findings and confirm the predominant location of the amine in the nervous tissue. These results suggest that histamine is more widespread than some other putative transmitters, and support the concept that histamine may have an important role in many physiological processes in molluscs.

Animals↗
PubMed1995

In vitro hormonal regulation of glycogen phosphorylase activity in fat body of the tropical cockroach, Blaberus discoidalis.

Blaberus hypertrehalosemic hormone (Bld-HTH)-dependent glycogen phosphorylase activation was investigated using in vitro fat bodies from the cockroach, Blaberus discoidalis. Resting levels of active phosphorylase were decreased by the presence of trehalose and glucose. Phosphorylase activation was dose-responsive to Bld-HTH and increased ca. 3-fold over a range of 0.02 to 2 nM Bld-HTH. Maximum phosphorylase activation required only 5-min exposure to Bld-HTH; reversion to the inactive state began within 15 min after Bld-HTH removal and was completed by 60 min. Octopamine also activated phosphorylase but required 10(3)-fold higher concentrations than did Bld-HTH. Concentrations of Bld-HTH and octopamine that increased active phosphorylase did not elevate fat body cAMP levels, although a high concentration of octopamine increased tissue cAMP levels. cAMP did not increase phosphorylase activity, but Ca2+ was important for both Bld-HTH- and octopamine-dependent phosphorylase activation.

Animals↗
PubMed1988

Efferent neurotransmission of circadian rhythms in Limulus lateral eye. II. Intracellular recordings in vitro.

We investigated efferent neurotransmission in the Limulus lateral eye by studying the action of pharmacological agents on responses of photoreceptor cells in vitro. We recorded transmembrane potentials from single cells in slices of retina that were excised during the day and maintained for several days in a culture medium. Potentials recorded in the absence of pharmacological agents resemble those recorded from cells in vivo during the day. Octopamine, a putative efferent neurotransmitter, induced changes in photoreceptor potentials that mimicked in part those generated at night by a circadian clock located in the brain. Specifically, octopamine (100 to 500 microM) decreased the frequency of occurrence of quantum bumps in the dark and increased the amplitude of photoreceptor responses to intermediate and high light intensities. Similar actions were produced by naphazoline (25 to 100 microM, potent agonist of octopamine), forskolin (8 to 400 microM, activator of adenylate cyclase), IBMX (1 mM, inhibitor of phosphodiesterase), and 8-bromo-cAMP (500 microM, analogue of cAMP). 8-bromo-cGMP (500 microM, analogue of cGMP) decreased the rate of spontaneous quantum bumps only. Our results support the hypothesis that (1) octopamine is an efferent neurotransmitter of circadian rhythms in the Limulus eye and that (2) it activates adenylate cyclase to increase levels of the second messenger, cAMP, in photoreceptor cells. Circadian changes in photoreceptor responses to moderate intensities may be a specific action of cAMP, since cGMP has no effect. Circadian changes in the rate of spontaneous quantum bumps may involve a less specific intermediate, since both cAMP and cGMP reduce bump rate. Characteristics of the retinal slice preparation precluded a detailed study of the effects of pharmacological agents on retinal morphology.

1-Methyl-3-isobutylxanthine↗
PubMed2003

Effect of pheromones, hormones, and handling on sucrose response thresholds of honey bees (Apis mellifera L.).

The responsiveness of bees to sucrose is an important indicator of honey bee foraging decisions. Correlated with sucrose responsiveness is forage choice behavior, age of first foraging, and conditioned learning response. Pheromones and hormones are significant components in social insect systems associated with the regulation of colony-level and individual foraging behavior. Bees were treated to different exposure regimes of queen and brood pheromones and their sucrose responsiveness measured. Bees reared with queen or brood pheromone were less responsive than controls. Our results suggest responsiveness to sucrose is a physiologically, neuronally mediated response. Orally administered octopamine significantly reduced sucrose response thresholds. Change in response to octopamine was on a time scale of minutes. The greatest separation between octopamine treated and control bees occurred 30 min after feeding. There was no significant sucrose response difference to doses ranging from 0.2 mug to 20 mug of octopamine. Topically applied methoprene significantly increased sucrose responsiveness. Handling method significantly affected sucrose responsiveness. Bees that were anesthetized by chilling or CO(2) treatment were significantly more responsive than control bees 30 min after handling. Sixty minutes after handling there were no significant treatment differences. We concluded that putative stress effects of handling were blocked by anesthetic.

Anesthesia↗
PubMed2001

Phenolamine-dependent adenylyl cyclase activation in Drosophila Schneider 2 cells.

Drosophila Schneider 2 (S2) cells are often employed as host cells for non-lytic, stable expression and functional characterization of mammalian and insect G-protein-coupled receptors (GPCRs), such as biogenic amine receptors. In order to avoid cross-reactions, it is extremely important to know which endogenous receptors are already present in the non-transfected S2 cells. Therefore, we analyzed cellular levels of cyclic AMP and Ca2+, important second messengers for intracellular signal transduction via GPCRs, in response to a variety of naturally occurring biogenic amines, such as octopamine, tyramine, serotonin, histamine, dopamine and melatonin. None of these amines (up to 0.1 mM) was able to reduce forskolin-stimulated cyclic AMP production in S2 cells. Furthermore, no agonist-induced calcium responses were observed. Nevertheless, the phenolamines octopamine (OA) and tyramine (TA) induced a dose-dependent increase of cyclic adenosine monophosphate (AMP) production in S2 cells, while serotonin, histamine, dopamine and melatonin (up to 0.1 mM) did not. The pharmacology of this response was similar to that of the octopamine-2 (OA2) receptor type. In addition, this paper provides evidence for the presence of an endogenous mRNA encoding an octopamine receptor type in these cells, which is identical or very similar to OAMB. This receptor was previously shown to be positively coupled to adenylyl cyclase.

Adenosine Monophosphate↗
PubMed2005

Mode of binding of methyl acceptor substrates to the adrenaline-synthesizing enzyme phenylethanolamine N-methyltransferase: implications for catalysis.

Here we report three crystal structure complexes of human phenylethanolamine N-methyltransferase (PNMT), one bound with a substrate that incorporates a flexible ethanolamine side chain (p-octopamine), a second bound with a semirigid analogue substrate [cis-(1R,2S)-2-amino-1-tetralol, cis-(1R,2S)-AT], and a third with trans-(1S,2S)-2-amino-1-tetralol [trans-(1S,2S)-AT] that acts as an inhibitor of PNMT rather than a substrate. A water-mediated interaction between the critical beta-hydroxyl of the flexible ethanolamine group of p-octopamine and an acidic residue, Asp267, is likely to play a key role in positioning the side chain correctly for methylation to occur at the amine. A second interaction with Glu219 may play a lesser role. Catalysis likely occurs via deprotonation of the amine through the action of Glu185; mutation of this residue significantly reduced the kcat without affecting the Km. The mode of binding of cis-(1R,2S)-AT supports the notion that this substrate is a conformationally restrained analogue of flexible PNMT substrates, in that it forms interactions with the enzyme similar to those observed for p-octopamine. By contrast, trans-(1S,2S)-AT, an inhibitor rather than a substrate, binds in an orientation that is flipped by 180 degrees compared with cis-(1R,2S)-AT. A consequence of this flipped binding mode is that the interactions between the hydroxyl and Asp267 and Glu219 are lost. However, the amines of inhibitor trans-(1S,2S)-AT and substrate cis-(1R,2S)-AT are both within methyl transfer distance of the cofactor. These results suggest that PNMT catalyzes transfer of methyl to ligand amines only when "anchor" interactions, such as those identified for the beta-hydroxyls of p-octopamine and cis-AT, are present.

Amino Acid Substitution↗
PubMed1987

Amphetamine and reserpine deplete brain biogenic amines and alter blow fly feeding behavior.

HPLC with electrochemical detection was used to determine the levels of p-hydroxyphenylethanolamine (octopamine), 3,4-dihydroxyphenylethylamine (dopamine), and 5-hydroxytryptamine (5-HT) in the brains of control, reserpine, and d-amphetamine-treated blow flies, Phormia regina Meigen. Parallel studies were carried out to assess the effects of the two drugs on fly feeding behavior, measured as mean acceptance threshold: the minimum sucrose concentration to which the average fly in a population will respond by proboscis extension when its tarsi contact the solution. In saline-injected control flies, all three amines were found at levels of approximately 2 pmol/brain. Thirty minutes after injection with d-amphetamine (12 micrograms/fly), brain octopamine was depleted by 85%, whereas dopamine and 5-HT were depleted by 70%. Reserpine (5 micrograms/fly) caused 70% depletion of dopamine and greater than 90% depletion of both octopamine and 5-HT 24 h after injection. However, the effect of reserpine was much slower in onset (hours versus minutes) and more persistent (days versus hours) than was the effect of d-amphetamine. With either drug, the time course of amine depletion closely matched the time course of the increase in feeding threshold observed in drug-treated flies. These results suggest that CNS pools of the biogenic amines, octopamine, dopamine, and 5-HT are important in governing blow fly responsiveness to food stimuli.

Amines↗
PubMed1989

Prenatal ontogenesis of brain phenolamines and catecholamines in relation to their metabolizing enzymes in Roman avoider strains of rats.

Phenolamines, particularly octopamines, are of special importance in avoidance behavior. In the Roman low avoidance (RLA) strain, p-octopamine can induce locomotor behavioral activity that is normally observed in the Roman high avoidance (RHA) strain. For these reasons, the levels of prenatal octopamines (para and meta isomers) have been studied in relation to noradrenaline and dopamine levels. In the hypothalamus and brainstem of RHA, a maximum level of the para isomer is observed at 15 days of embryonic development but, unlike in controls and RLA animals, this level remains almost constant until 20 days. For the meta-isomer and catecholamines, there is a 1-2 day delay in detection between controls and RLA or RHA. The study of related enzyme activities reveals that tyrosine hydroxylase displays a 2-day delay in RHA when compared to the control value at 19 days of fetal life. These results are discussed in terms of the role of p-octopamine in avoidance conditioning and of the possible delayed expression of the tyrosine hydroxylase gene in Roman strains of rats.

2-Hydroxyphenethylamine↗