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Phylogenetic distribution of aromatase and other androgen-converting enzymes in the central nervous system.

Metabolism of [3H]androstenedione was studied in brain tissue homogenates of opossum, bird, snake, sea turtle, urodele amphibian, teleost, shark, skate, hagfish, and lobster. Estrone, 17 beta-estradiol, or 17 alpha-estradiol was formed by central neural tissues of all species, with the exception of the opossum, hagfish, and lobster. Aromatase activity was concentrated in the forebrain, although some estrogen was synthesized by mid- or hindbrain homogenates of two lower vertebrates (teleost and skate) and the newly hatched chick. 5 alpha-Androstanedione (5 alpha-androstane-3,17-dione) or 5 alpha-dihydrotestosterone were products of metabolism in several nonmammalian vertebrates and in the invertebrate central nervous system also. 5 alpha-Reductase was found in all major brain divisions. These and previously reported comparative studies indicate that the ability to aromatize and otherwise transform androgen substrates is a primitive characteristic of the brain that has been widely conserved phylogenetically.

Androstenedione↗

Locomotion by Abdopus aculeatus (Cephalopoda: Octopodidae): walking the line between primary and secondary defenses.

Speeds and variation in body form during crawling, bipedal walking, swimming and jetting by the shallow-water octopus Abdopus aculeatus were compared to explore possible interactions between defense behaviors and biomechanics of these multi-limbed organisms. General body postures and patterns were more complex and varied during the slow mode of crawling than during fast escape maneuvers such as swimming and jetting. These results may reflect a trade-off between predator deception and speed, or simply a need to reduce drag during jet-propelled locomotion. Octopuses swam faster when dorsoventrally compressed, a form that may generate lift, than when swimming in the head-raised posture. Bipedal locomotion proceeded as fast as swimming and can be considered a form of fast escape (secondary defense) that also incorporates elements of crypsis and polyphenism (primary defenses). Body postures during walking suggested the use of both static and dynamic stability. Absolute speed was not correlated with body mass in any mode. Based on these findings the implications for defense behaviors such as escape from predation, aggression, and 'flatfish mimicry' performed by A. aculeatus and other octopuses are discussed.

Animals↗

Electromyography of the buccal musculature of octopus (Octopus bimaculoides): a test of the function of the muscle articulation in support and movement.

The buccal mass musculature of the octopus (Octopus bimaculoides) was studied with electromyography to test the predictions of a previous morphological study in which we suggested that the muscles of the buccal mass serve as both the effectors of movement and as the joint itself, forming a new category of flexible joint termed a ;muscle articulation'. The predictions of muscle function were tested by correlating muscle electrical activity in isolated buccal masses with spontaneous beak movements. Bipolar electromyography electrodes were implanted in the various beak muscles and beak position was recorded simultaneously with an electronic movement monitor (N=14). The results are consistent with the hypothesis that the lateral mandibular muscles produce opening movements of the beaks and provide the first definitive explanation of the opening mechanism. The results are also consistent with the hypothesis that the superior mandibular muscle functions primarily in closing. Co-contraction of the lateral mandibular muscles and the superior mandibular muscles was also observed, suggesting that these muscles may also stabilize the beaks during movement or provide a means of controlling the location of the pivot between the beaks. This study provides an important first test of the predictions of the role of the complex musculature found in muscle articulations such as the cephalopod buccal mass.

Animals↗

Strontium is required for statolith development and thus normal swimming behaviour of hatchling cephalopods.

When cephalopod eggs were incubated in artificial sea water it was found that they sometimes resulted in hatchlings with defects of the statocyst suprastructure, leading to the severe behavioural defect of uncontrolled swimming. Experiments in defined media (seven basic salts mixed in deionized water) with seven species of cephalopods demonstrated clearly that there is 100% normal development of the aragonite statoliths when strontium levels were 8 mg l-1. Conversely, statoliths did not develop when strontium was absent. In cuttlefish, the growth of the cuttlebone was also affected adversely when strontium was absent. In mariculture production tanks, supplementing commercial artificial sea water with strontium to normal levels of 8 mg l-1 almost eliminated the occurrence of abnormal hatchlings. Circumstantial evidence indicates that there is a critical window in development during which strontium is required for normal development. The role of strontium in biomineralization during embryogenesis is unknown, but it appears to be important in the Mollusca.

Animals↗

Directional sensitivity of hair cell afferents in the Octopus statocyst.

Changes in threshold sensitivity of hair cell afferents of the macula and crista of the Octopus statocyst were analyzed when the hair cells were stimulated with sinusoidal water movements from different directions. The experiments indicate that cephalopod statocyst hair cells are directionally sensitive in a way that is similar to the responses of the hair cells of the vertebrate vestibular and lateral line systems, with the amplitude of the response changing according to the cosine of the angle by which the direction of the stimulus (the deflection of the ciliary bundle) deviates from the direction of the hair cell's morphological polarization.

Action Potentials↗

Polarization contrast vision in Octopus.

While the ability to analyze polarized light is widespread among animals, its contribution to form vision has not yet been documented. We tested the hypothesis that polarization vision can be used for object discrimination, by training octopuses to distinguish between targets on the basis of the presence or absence of a pattern produced by a 90 degrees polarization contrast within the target. Octopuses recognized a 90 degrees contrast pattern within a single target, when presented either on a horizontal/vertical axis or on a 45 degrees/135 degrees axis. They were able to transfer their learning to new situations and to detect a polarization contrast when the orientations of the e-vector of light passing through the target center and background differed by as little as 20 degrees. Polarization vision may provide information similar to that available from color vision and thus serve to enhance the detection and recognition of objects.

Animals↗

An in vitro bioassay for a mulluscan gonadotropin.

1. Protein synthesis occurs at a high rate in the ovaries of maturing Octopus vulgaris and can be measured from the incorporation of [14C]leucine in vivo and in isolated groups of eggs in vitro. 2. Removal of the optic glands in vivo 1--3 days prior to testing markedly reduces amino acid incorporation in vivo or in vitro. After 5 days in vivo incorporation stops. 3. The rate of incorporation in vitro is increased by the addition of optic gland extract. 4. Analysis of the kinetics of leucine uptake and incorporation in vitro indicates that the hormone has an effect on the inward transport of leucine which is independent of its action on protein synthesis. 5. Electron-microscope studies of the follicle cells and ova show that the former are the site of protein synthesis. 6. Changes in either uptake or incorporation into protein by the follicle cells can be used as a qualitative biolobical assay for the optic gland hormone. Uptake is very easy to measure but incorporation is the more sensitive parameter. Either is potentially suitable as a quantitative assay for this and perhaps also for other molluscan gonadotropins.

Animals↗

Optic gland implants and their effects on the gonads of Octopus.

1. Optic glands transplanted from one Octopus vulgaris into another cause enlargement of the gonads and ducts of the recipient. 2. Enlargement occurs whether or not the gland was secreting when implanted and regardless of the sex of the donor or recipient. 3. Glands derived from Eledone moschata or Octopus macropus implanted into O. vulgaris are as effective as glands derived from O. vulgaris. 4. Implants derived from Sepia officinalis or Loligo vulgaris appear to be ineffective.

Animals↗

Receptor units responding to movement in the octopus mantle.

1. A preparation of the mantle of Octopus which is inverted over a solid support and which exposes the stellate ganglion and associated nerves is described. 2. Afferent activity can be recorded from stellar nerves following electrical stimulation of the pallial nerve. The latency and frequency of the phasic sensory response is correlated with the contraction of the mantle musculature. 3. It is proposed that receptors cells located in the muscle, and their activity following mantle contraction, form part of a sensory feedback system in the mantle. Large, multipolar nerve cells that were found between the two main layers of circular muscle in the mantle could be such receptors.

Action Potentials↗

Reproduction versus somatic growth: hormonal control in Octopus vulgaris.

1. Octopus vulgaris can be forced into precocious maturity by removal of the subpedunculate lobe from the brain, an operation that releases the optic glands from inhibition, and allows them to secrete a gonadotropin. 2. 14C-leucine was injected into the bloodstream of immature animals and its subsequent incorporation into muscle protein followed by taking successive samples from the arms. The optic glands were then activated, and a further injection of 3H-leucine given and followed by means of further arm samples. 3. Optic gland secretion suppresses protein synthesis in the muscles. This is associated with an increase in the total amino acid pool in the muscles and with a considerable increase in the concentration of free amino acids circulating in the blood. 4. If an ovary is present these events are associated with a rapid growth of the ovary and its ducts, and a loss of weight elsewhere. In ovariectomized animals the ducts grow, but there is no yolk to absorb the large pool of free amino acids, and the animals gain weight by osmotic uptake of water into the muscles. 5. The developing ovary may produce a hormone that increases the release of amino acids from muscle, since the concentration circulating in the blood of intact animals remains at least as high as in ovariectomized octopuses, despite the demands of the developing ovary. 6. These matters are discussed in relation to other evidence for a gonadial hormone and in relation to the 'self-destruct' effect of the optic gland secretion in determining the post-reproductive death of octopuses.

Age Factors↗

The effects of extracts from neurosecretory cells in the anterior vena cava and pharyngo-ophthalmic vein upon the hearts of intact free-moving octopuses.

Recordings of pressure and frequency were made from the hearts of free-moving Octopus vulgaris. The effects of extracts from neurosecretory endings in the anterior vena cava (AVC) and the pharyngo-ophthalmic vein (POV), injected through fine cannulae into a branchial heart, efferent branchial vessel or the dorsal aorta, were studied and compared with the effects of acetylcholine, 5-hydroxytryptamine, adrenaline, histamine and tyramine. AVC and POV extracts each produce a different spectrum of effects, unlike those of any of the drugs tested. AVC extract is effective at doses of less than 2% of the material extractable from a single vein per kg, increasing the force and amplitude of the heartbeats. With a natural release point just upstream of the branchial hearts the AVC material must be relevant to the normal performance of the hearts. POV extract is effective only at doses equivalent to several veins per kg, and is unlikely to have a role in cardiac regulation. Section of the visceral nerves did not affect the action of drugs or extracts, indicating that effects were not indirectly mediated via the CNA. Further experiments were made with hearts and the aorta in vitro with effects that did not always parallel those found in vivo. Reasons for these differences are discussed.

Acetylcholine↗

Nervous control of the heartbeat in octopus.

The circulatory system of cephalopods is based on a trio of hearts, with two pairs of associated ganglia linked to the CNS by a pair of visceral nerves. The beat of the hearts was recorded from free-moving octopuses before and after surgical removal or disconnexion of elements of the nervous system. Severing the visceral nerves does not stop the hearts, which continue to beat in a powerful well co-ordinated manner in isolation from the CNS. The nerves seem to be concerned in raising the cardiac output in exercise, and with stopping the hearts when mantle movements cease, but they are not necessary for the initiation of maintenance of the normal rhythm. Removal of the fusiform ganglia severs all nervous connexions between the ywo gill hearts, and deprives the systemic heart of its nerve supply. The trio of hearts continues to beat as strongly as before. Removal or disconnexion of a cardiac ganglion disrupts the beat of the corresponding gill heart which now tends to contract in an ill-coordinated and rather feeble manner, though at much the same frequency as before; with both cardiacs gone the systemic heart, which contracts only when it is filled, tends to drop in frequency and the mean aortic pressure falls. The system remains rhythmic, however, and the beat may recover, to the point where aortic pressures and frequencies approach those found in intact animals at rest; even octopuses with both fusiform and both cardiac ganglia removed can survive for many hours. From the performance of the isolated branchial heart, the existence of a pulsating vesicle in each cardiac ganglion, the effects of cardiac ganglion removal and the remarkable steadiness of heartbeat frequency shown by intact animals under a variety of conditions, it is argued that the heartbeat rhythm is normally controlled by pacemakers in the branchial heart/ cardiac ganglion complexes, and perhaps, in intact animals, from within the cardiac ganglia themselves. The picture of the control of the heartbeat that emerges from the study of free moving essentially intact animals is quite different from that arising from in vitro and acute preparation studies. It suggests that the conventional wisdom about the control of the heartbeat in cephalopods (and perhaps by implication, in other molluscs) may need to be considerably revised.

Animals↗