The active transport of oxygen and carbon dioxide into the swim-bladder of fish

The active transport of oxygen and carbon dioxide into the swim-bladder of fish is discussed. The rete mirabile is a capillary network which is involved in the gas secretion into the bladder. The rete is regarded as a counter-current multiplier. Lactic acid which is produced in the gas gland generates in the rete single concentrating effects for oxygen and carbon dioxide; i.e., for equal partial pressures the concentrations of the gases in the afferent rete capillaries are higher than those in the efferent ones. The single concentrating effects were calculated from measurements of sea robin blood (Root, 1931). The multiplication of these effects within the rete for different rete lengths and different transport rates was numerically evaluated. The calculated O₂ and CO₂ pressures in the bladder are in good agreement with the experimental results of Scholander and van Dam (1953). The descent velocities at equilibrium between bladder pressure and hydrostatic pressure are discussed for fishes with different rete lengths.     

The Secretion of Oxygen into the Swim-bladder of Fish : II. The simultaneous transport of carbon monoxide and oxygen

Toadfish, Opsanus tau, L., were maintained in sea water equilibrated with gas mixtures containing a fixed proportion of oxygen and varying proportions of carbon monoxide. The swim-bladder was emptied by puncture, and, after an interval of 24 or 48 hours, the newly secreted gases were withdrawn and analyzed. Both carbon monoxide and oxygen are accumulated in the swim-bladder at tensions greater than ambient. The ratio of concentrations, carbon monoxide (secreted): carbon monoxide (administered) bears a constant relation to the ratio, oxygen (secreted): oxygen (administered). The value of the partition coefficient describing this relation is (α = 5.44). The two gases are considered to compete for a common intracellular carrier mediating their active transport. The suggestion is advanced that the intracellular oxygen carrier is a hemoglobin. Comparison of the proportions of carboxy- and oxyhemoglobin in the blood with the composition of the secreted gas proves that the secreted gases are not evolved directly from combination with blood hemoglobin. The suggestion is advanced that cellular oxygen secretion occurs in the rete mirabile: the rete may build up large oxygen tensions in the gas gland capillaries. It is suggested that the gas gland acts as a valve impeding back diffusion of gases from the swim-bladder.     

O2-filled swimbladder employs monocarboxylate transporters for the generation of O2 by lactate-induced root effect hemoglobin

The swimbladder volume is regulated by O(2) transfer between the luminal space and the blood In the swimbladder, lactic acid generation by anaerobic glycolysis in the gas gland epithelial cells and its recycling through the rete mirabile bundles of countercurrent capillaries are essential for local blood acidification and oxygen liberation from hemoglobin by the "Root effect." While O(2) generation is critical for fish flotation, the molecular mechanism of the secretion and recycling of lactic acid in this critical process is not clear. To clarify molecules that are involved in the blood acidification and visualize the route of lactic acid movement, we analyzed the expression of 17 members of the H(+)/monocarboxylate transporter (MCT) family in the fugu genome and found that only MCT1b and MCT4b are highly expressed in the fugu swimbladder. Electrophysiological analyses demonstrated that MCT1b is a high-affinity lactate transporter whereas MCT4b is a low-affinity/high-conductance lactate transporter. Immunohistochemistry demonstrated that (i) MCT4b expresses in gas gland cells together with the glycolytic enzyme GAPDH at high level and mediate lactic acid secretion by gas gland cells, and (ii) MCT1b expresses in arterial, but not venous, capillary endothelial cells in rete mirabile and mediates recycling of lactic acid in the rete mirabile by solute-specific transcellular transport. These results clarified the mechanism of the blood acidification in the swimbladder by spatially organized two lactic acid transporters MCT4b and MCT1b.     

Limits of gas secretion by the salting-out effect in the fish swimbladder rete

Summary The high dissolved gas tensions required for the secretion of gases into deep-sea fish swimbladders are thought to be produced in the rete mirabile by a countercurrent multiplication mechanism, the capacity of which is theoretically limited by the physical characteristics of the rete and by the magnitudes of minute gas solubility changes in the blood plasma. These gas solubility changes are presumably induced through the salting-out effect following the addition of lactic acid to rete venous blood as it circulates through the gas gland. In order to estimate the maximum swimbladder gas pressures attainable by this mechanism, the effects of lactic acid on N₂ and Ar solubilities in water were determined at 5 and 25°C with a new volumetric method. The results show that the salting-out effect with lactic acid is much smaller than with NaCl, and that the agreement between predicted and observed swimbladder gas pressures is more critically dependent on the physical properties of the rete vasculature than indicated by previous theoretical treatments. When augmented by the release of hemoglobin-bound O₂, the salting-out effect with lactic acid appears large enough to account for the production of even the highest swimbladder O₂ pressures, provided the rete characteristics lie within certain reasonable limits. However, successful theoretical explanation of observed swimbladder N₂ pressures in some deep-sea species will require rigorous attention to such theoretically neglected factors as dissolved gas backdiffusion along the rete and the unequal size and number of the rete arterial and venous capillaries.     

Comparison of cerebral base arteries in antelopes of Tragelaphus,Taurotragus and Boselaphus genera

Studies were conducted on preparations of head arteries, including cerebral base arteries of sitatunga (n = 14), nyala n = 16) and Greater kudu n = 4) of Tragelaphus genus, Common eland n = 7) from the tribe of Tragelaphini and the largest Asiatic Nilgai antelope n = 9) from Boselaphini tribe. Blood vessels of the cerebral arterial circle in studied antelopes were found to arise from terminal division of intracranial segments of internal carotid arteries, which emerge from the pairwise rostral epidural rete mirabile. Due to obliteration of the extracranial segment of internal carotid artery, the cerebral arterial circle of studied antelopes is supplied with blood mainly through maxillary artery, with mediation of blood vessels of rostral epidural rete mirabile. The system of cerebral base arteries in selected representatives of Tragelaphus, Taurotragus and Boselaphus, in contrast to other species of Bovinae subfamily, manifests the absence of caudal epidural rete mirabile. The pattern of cerebral base arteries in studied antelopes is consistent with position of the species in taxonomy worked out by Simpson (Bull Am Mus Nat Hist 85:1–350, 1945) and McKenna and Bell (Classification of mammals above the species level. Columbia University Press, New York, pp I–XII, 1997).     

Structure of the Rete Mirabile in the Kidney of the Rat as Seen with the Electron Microscope

Electron micrographs of the rete mirabile in the medulla of the rat have revealed that the endothelium of the afferent and efferent vessels are markedly different in fine structure. The venous capillaries returning blood from the papilla are lined with a fenestrated endothelium much like that in the peritubular capillaries of the kidney. The arterial capillaries delivering blood to the papilla have an unperforated lining of overlapping endothelial cells with extremely irregular tapered margins. It is pointed out that the organization of particularly the latter vessels suggests that the functional capabilities of these retia go beyond those of a simple diffusion countercurrent exchanger.     

Short Circuiting of the Ocular Oxygen Concentrating Mechanism in the Teleost Salmo gairdneri Using Carbonic Anhydrase Inhibitors

Ocular oxygen concentration by the process of counter current multiplication in rainbow trout (Salmo gairdneri) was rapidly suppressed after intraperitoneal injections of the carbonic anhydrase inhibitor CL-11,366. The rapidity with which this drug acted suggested a short circuiting of the choroidal rete mirabile. A comparison was made between the time after injection of inhibitor at which oxygen concentrating ability was lost to the time after injection of inhibitor at which its presence in red blood cells, choroidal rete, pseudobranch, and retinal tissue was first noted. A scheme for the possible role of carbonic anhydrase from each of these tissues in the process of ocular oxygen concentration is given.     

The secretion of inert gas into the swim-bladder of fish

The composition of the gas mixture secreted into the swim-bladders of several species of fish has been determined in the mass spectrometer. The secreted gas differed greatly from the gas mixture breathed by the fish in the relative proportions of the chemically inert gases, argon, neon, helium, and nitrogen. Relative to nitrogen the proportion of the very soluble argon was increased and the proportions of the much less soluble neon and helium decreased. The composition of the secreted gas approaches the composition of the gas mixture dissolved in the tissue fluid. A theory of inert gas secretion is proposed. It is suggested that oxygen gas is actively secreted and evolved in the form of minute bubbles, that inert gases diffuse into these bubbles, and that the bubbles are passed into the swim-bladder carrying with them inert gases. Coupled to a preferential reabsorption of oxygen from the swim-bladder this mechanism can achieve high tensions of inert gas in the swim-bladder. The accumulation of nearly pure nitrogen in the swim-bladder of goldfish (Carassius auratus) is accomplished by the secretion of an oxygen-rich gas mixture followed by the reabsorption of oxygen.     

Gas Exchange of Algae: II. Effects of Oxygen, Helium, and Argon on the Photosynthesis of Chlorella pyrenoidosa

Changes in the oxygen partial pressure of air over the range of 8 to 258 mm of Hg did not adversely affect the photosynthetic capacity of Chlorella pyrenoidosa. Gas exchange and growth measurements remained constant for 3-week periods and were similar to air controls (oxygen pressure of 160 mm of Hg). Oxygen partial pressures of 532 and 745 mm of Hg had an adverse effect on algal metabolism. Carbon dioxide consumption was 24% lower in the gas mixture containing oxygen at a pressure 532 mm of Hg than in the air control, and the growth rate was slightly reduced. Oxygen at a partial pressure of 745 mm of Hg decreased the photosynthetic rate 39% and the growth rate 37% over the corresponding rates in air. The lowered metabolic rates remained constant during 14 days of measurements, and the effect was reversible after this time. Substitution of helium or argon for the nitrogen in air had no effect on oxygen production, carbon dioxide consumption, or growth rate for 3-week periods. All measurements were made at a total pressure of 760 mm of Hg, and all gas mixtures were enriched with 2% carbon dioxide. Thus, the physiological functioning and reliability of a photosynthetic gas exchanger should not be adversely affected by: (i) oxygen partial pressures ranging from 8 to 258 mm of Hg; (ii) the use of pure oxygen at reduced total pressure (155 to 258 mm of Hg) unless pressure per se affects photosynthesis, or (iii) the inclusion of helium or argon in the gas environment (up to a partial pressure of 595 mm of Hg).     

Histological demonstration of glucose transporters,fructose-1,6-bisphosphatase,and glycogen in gas gland cells of the swimbladder: Is a metabolic futile cycle operating?

Luminal surface of the swimbladder is covered by gas gland epithelial cells and is responsible for inflating the swimbladder by generating O(2) from Root-effect hemoglobin that releases O(2) under acidic conditions. Acidification of blood is achieved by lactic acid secreted from gas gland cells, which are poor in mitochondria but rich in the glycolytic activity. The acidic conditions are locally maintained by a countercurrent capillary system called rete mirabile. To understand the regulation of anaerobic metabolism of glucose in the gas gland cells, we analyzed the glucose transporter expressed there and the fate of ATP generated by glycolysis. The latter is important because the ATP should be immediately consumed otherwise it strongly inhibits the glycolysis rendering the cells unable to produce lactic acid anymore. Expression analyses of glucose transporter (glut) genes in the swimbladder of fugu (Takifugu rubripes) by RT-PCR and in situ hybridization demonstrated that glut1a and glut6 are expressed in gas gland cells. Immunohistochemical analyses of metabolic enzymes demonstrated that a gluconeogenesis enzyme fructose-1,6-bisphosphatase (Fbp1) and a glycolytic enzyme glyceraldehyde-3-phosphate dehydrogenase (Gapdh) are highly expressed in gas gland cells. The simultaneous catalyses of glycolysis and gluconeogenesis reactions suggest the presence of a futile cycle in gas gland cells to maintain the levels of ATP low and to generate heat that helps reduce the solubility of O(2).     

The Dependence of the Oxygen-Concentrating Mechanism of the Teleost Eye (Salmo gairdneri) on the Enzyme Carbonic Anhydrase

Microoxygen polarographic electrodes were constructed and used to measure oxygen tension (P_OO2) in the eyes of rainbow trout (Salmo gairdneri). The values obtained are compared with arterial blood and environmental water P_OO2 and indicate that there is an oxygen-concentrating mechanism in the eye supplying oxygen to the avascular retina. Anatomically similar retes suggest that the mechanism is similar to the one which exists in the swim bladder. Elimination of the arterial blood supply to the choroidal gland rete mirabile of the eye (through pseudobranchectomy) and the consequent lowering of ocular oxygen tensions implicate the choroidal gland as one of the major components of the oxygen-concentrating mechanism. After pseudobranchectomy the presence of ocular P_OO2 above that of arterial blood is indicative of a secondary structure in the eye capable of concentrating oxygen. Inhibition of carbonic anhydrase, using acetazolamide, is shown to result in complete suppression of the oxygen-concentrating mechanism. A hypothesis is advanced for the participation of retinal-choroidal and erythrocyte carbonic anhydrase in the oxygen-concentrating mechanism.     

Structure and sympathetic innervation of the intracranial arteries in the giraffe (Giraffa camelopardalis)

Fluorescence histochemistry discloses that the carotid rete mirabile in the giraffe has a poor sympathetic innervation. In contrast, the efferent artery of the rete (internal carotid artery) and the cerebral arteries show moderate sympathetic innervation. A certain degree of regional variability was noted in which the rostral arteries (anterior and middle cerebral) receive more sympathetic nerves than the caudal (posterior communicating and basilar) arteries. The sympathetic nerves on the giraffe cerebral vessels may constitute part of a host of mechanisms by which regional blood flow to the brain is regulated. Conversely, the paucity of sympathetic innervation of the carotid rete mirabile may indicate that this structure does not play an active role in vasoconstrictor responses during postural changes of the head.     

Interactions between bluefish and striped bass: Behavior of bluefish under size- and number-impaired conditions and overlap in resource use

A decline in bluefish (Pomatomus saltatrix L.) recreational landings during the 1990s and the early 2000s led to multiple theories on the ultimate cause. One theory was that a large portion of the bluefish population moved offshore and was unavailable to nearshore recreational fishers; one reason given for the movement offshore was increased competition with striped bass (Morone saxatilis W.). We conducted laboratory experiments (feeding and non-feeding) to examine behavioral interactions between adult bluefish and sub-adult striped bass in a large (121,000 L) research aquarium. Additionally, we examined diet and habitat overlap of bluefish and striped bass from the fall and spring bottom trawl surveys conducted by the National Marine Fisheries Service. Observations of feeding trials for the following treatments were made: non-impaired (i.e., same number and size of bluefish and striped bass), size-impaired (i.e., large striped bass/small bluefish), number-impaired (i.e.,10 striped bass/3bluefish), and single-species controls. Within a species, there was no difference in a variety of behavioral measures (e.g., attack rate, capture success, ingestion rate, and activity) between mixed- and control treatments under non-impaired or size-impaired conditions. However, behavior of number-impaired bluefish differed from control and size-impaired fish suggesting that striped bass may have a negative influence on bluefish foraging when bluefish are “out-numbered”. Feeding had a strong effect on swimming speeds for both species. Diet and habitat overlap between bluefish and striped bass in continental shelf waters was low. Overall, foraging behavior in mixed-species treatments and field observations suggest no competitive interactions between adult bluefish and sub-adult striped bass.     

Der O2-Austausch zwischen Capillaren und Gewebe

Summary The distribution of oxygen concentration or oxygen partial pressure (=po₂) in models with different capillary arrangements is calculated. The capillaries form homogeneously perfused concurrent and countercurrent systems or inhomogeneously perfused capillary mesh systems with different velocities of blood flow on the inside of the capillaries. The question arises as to what extent the distribution of the po₂ in the tissue and the capillaries depends on the differences in streaming velocities and directions of the blood. The results show, that the greatest gradients in the po₂ distribution are to be found in the capillary mesh system. A typical example with a set of constants after Thews (1960) shows that the differences between the venous po₂ and the lowest po₂ in the tissue is 32 mm Hg in the case of the capillary network, 9 mm Hg in the case of the concurrent and 3 mm Hg in that of the countercurrent system.     

Oxidation of certain cyclic carbonyl compounds with alkaline hydrogen peroxide

The following cyclic compounds were oxidized at 4° by means of aqueous sodium peroxide in large excess: 2,3,4,6/5-pentahydroxycyclohexanone (1); xylo-pentahydroxy-2-cyclohexen-l-one (2); xylo-trihydroxycyclohexane-1,2,3-trione (3); reductic acid (4); and ninhydrin (5). On oxidation, 1, 2, and 3 gave carbon dioxide, formic acid, and oxalic acid; in addition, 1 yielded DL-glucaric acid, and 2 and 3 afforded xylaric acid. Compound 4 gave almost equimolecular quantities of carbon dioxide and succinic acid. In addition to a small amount of formic acid, compound 5 gave carbon dioxide and phthalic acid in yields of 92 and 95%, respectively. Reaction mechanisms are proposed.     

Cytochrome P4501A induced differentially in endothelial cells cultured from different organs of Anguilia rostrata

Summary Endothelial cells are a structural barrier and an active regulator of many bodily processes. Cytochrome P4501A (CYPIA) activity is induced in the endothelium of teleosts and mammals exposed to lipophilic xenobiotics, such as polycyclic aromatic hydrocarbons, and can have significant consequences for endothelial functions. We exposed cultures of characterized endothelial cells from the heart, kidney, and rete mirabile of the eel, Anguilla rostrata, to aryl hydrocarbon receptor (AhR) agonists. In heart endothelial cells, the maximum response (based on O-deethylation of 7-ethoxyresorufin to resorufin [EROD] activity) to 2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD), 113 pmol/mg/min, was at 1 nM TCDD and the peak response to β-napthoflavone (βNF), 135 pmol/mg/min, was at 3 μM βNF. The maximum response to TCDD in the kidney endothelial cells is 12 pmol/mg/min at 0.3 nM TCDD. The rete mirabile capillary endothelial cells responded minimally or not at all to exposure to TCDD and βNF. Both the heart and kidney endothelial cells (but not the rete mirabile capillary cells) have a low level of EROD activity (12.7 and 5.2 pmol/mg/min, respectively) in untreated or dimethylsulfoxide-treated cells. The robust response of the heart endothelial cells to induction and the lack of response in the rete mirabile capillary endothelial cells indicate that these cells are a good resource to use to investigate the physiological consequences of AhR agonist exposure and CYP1A induction in different areas of the vasculature.     

Evaluation of Hyperventilation in Treatment of Head Injuries

Reduction of the partial pressure of carbon dioxide in the arterial blood by mechanical hyperventilation (Pco₂ 25-30 mm Hg; Po₂ 100-150 mm Hg) may be beneficial in cases of severe head injury. To evaluate its efficacy and establish prognostic guidelines intracranial pressure, radiocirculograms, and cerebrospinal fluid (C.S.F.) lactate levels were studied in 31 patients. In survivors intracranial pressure fell and cerebral blood flow improved with treatment. A C.S.F. lactate greater than 55 mg/100 ml was associated with a poor prognosis. Selection of patients was based on clinical judgement, and adults with signs of extensive brain damage were excluded. The importance of an adequate airway and resuscitation is stressed before a final decision is made. The object of treatment is to improve the quality of survival and the criteria measured may aid in the distinction between patients with a potential for good recovery and those capable only of a vegetative existence. Many associated factors as well as hypocapnia reduce intracranial pressure, and these are discussed. We believe that hyperventilation may improve some head injuries, and further study is indicated.     

Microvasculature of the avian eye: studies on the eye of the duckling with microcorrosion casting, scanning electron microscopy, and stereology

The microvasculature of the eye of the duckling was studied with microcorrosion casting, scanning electron microscopy, and stereology. Most blood to the eyeball first passes through the arterial ophthalmic rete mirabile, a complex of small arteries which intermixes with a similar complex of veins (venous ophthalmic rete mirabile) at the ventrotemporal angle of the eye. The present study reveals the ultrastructural anatomy and the compact, three-dimensional arrangement of vessels in this rete, which had been shown by previous investigators to function as a countercurrent heat exchanger. Vessels from this rete include the supraorbital and infraorbital arteries, which supply the eyeball anteriorly, and the ophthalmotemporal artery, which supplies the eyeball posteriorly. The internal ophthalmic and ethmoidal arteries, branches of the cerebral carotid artery, anastomose with the ophthalmotemporal artery posteriorly. Blood is distributed to the eyeball anteriorly by two ring arteries: the iridial ring artery, which circumscribes the iris and which receives blood from the long ciliary and infraorbital arteries; and the more peripheral, ciliary ring artery, which receives blood mostly from the infraorbital and ethmoidal arteries. Within the iris is a dense, freely anastomosing bed of capillaries which extends to the edge of the pupil and then loops back beneath the ciliary body. The vasculature of the ciliary body consists of radially arranged plates of anastomosing capillaries of irregular bore which mimic the contours of that organ, but permit changes in pupil diameter. The present study demonstrates the three-dimensional anatomy of the very dense capillary net of the choriocapillaris deep to the retina and the capillary mass of the pecten, and thus supports the finding of earlier investigators that nutrients diffusing from these structures nourish the avascular retina. The pecten consists of a pleated sheet of freely anastomosing capillaries which protrudes into the vitreous body from near the optic nerve. The choriocapillaris and the pecten are supplied by branches of the ophthalmotemporal artery: the former by numerous short posterior ciliary arteries, the latter by two or three arteries which further divide into one or two smaller vessels for each of its folds. Veins of the choroid layer at the periphery of the anterior surface of the eyeball, and to some extent on its lateral walls, are revealed by the corrosion-casting technique as unusual, flattened vessels of large caliber which lie in closely spaced parallel arrays. The large surface area thus created may function in heat dissipation.(ABSTRACT TRUNCATED AT 400 WORDS)     

Formation of cyanate and carbamyl phosphate by electric discharges of model primitive gas

A mixed gas of nitrogen, carbon dioxide, and hydrogen was discharged over 100 ml of 0.2M NaHCO₃ solution in a 5 liter discharge apparatus which simulates the primitive Earth. The formation of cyanate, which is one of the possible primitive condensing agents, was demonstrated by the detection of [Cu(Py)₂] (NCO)₂ that was formed by the addition of copper sulfate-pyridine reagent to the solution. In a series of experiments the partial pressures of nitrogen and carbon dioxide in the starting gas were fixed at 10 cm Hg and 20 cm Hg, respectively, whereas that of hydrogen was varied between 5, 10, 15, 20, 25, and 30 cm Hg. The discharges were continued for one week. The rate of appearance of cyanate was strongly dependent upon the partial pressure of hydrogen. The maximum rate of the production of cyanate at the initial stage of the discharge was in the case of 10 cm Hg of hydrogen, in which condition the starting gas is in a predominantly oxidized state. In this case the concentration of cyanate reached about 0.012M after one day. Another discharge experiment was carried out with 0.2M phosphate solution, and the production of carbamyl phosphate was demonstrated through the formation of ATP by the incubation of the discharged solution with ADP and carbamyl phosphokinase.