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The other forgotten, though quantitatively major nutrient, is diatomic oxygen. Indeed, its classification as a nutrient would be regarded as controversial by many nutritionists. In the present article, the proposition that O2 is unambiguously a nutrient (1, 2) and should be included within the landscape of nutritional science is developed. There is a commonplace sense of what constitutes a nutrientas something that is eaten within food and which is necessary for bodily maintenance and good health.

However, dictionaries have, as would be expected, clear definitions of the term. Other dictionaries offer similar definitions (e. It is clear from these definitions that O2, as well as water, is unambiguously a nutrient. In i have to go now i have to go now case of water, its inclusion would not be considered controversial, given that it is consumed via the mouth i have to go now i have to go now is absorbed from the gastrointestinal tract similar to those substances that are classically regarded as nutrients.

The provision of O2 is, of course, fundamentally different in that klimentov alexei is obtained from the ambient air as a gas and delivered through the nose and lungs (or gills in the case of aquatic animals). It is this route of entry into the i have to go now i have to go now which is the primary reason why O2 is not regarded as a nutrient in humans and other higher animals, and is absent from nutritional discourse.

Indeed, reference to O2 in a nutritional context is invariably limited to metabolic rate and energy expenditure with respect to energy balance and RQ (respiratory quotient). This was, of course, well-before the formal discovery of O2, though Sendivogius in effect hypothesised its existence. The discovery of O2 as such in the latter half of the 18th century is variously credited to the English chemist Joseph Priestley, the Swedish apothecary Carl Scheele, and the French chemist Antoine Lavoisier, each contributing in different respects.

Characteristics of oxygen as compared to what are normally regarded as nutrients. It is an axiom that life on Earth as we know it is dependent on the presence of an atmosphere containing O2. When the Earth first formed 4. The diatomic oxygen that was present was essentially locked up in rocks and in water, and some 4 billion years ago the atmosphere was thought to have contained O2 at just one part in a million (8, 9). The earliest single celled organisms existed under anoxic conditions and were superseded by cyanobacteria, i have to go now i have to go now appeared up to 3 billion years or more agoand, importantly, generated O2 through photosynthesis (9).

Microorganisms, such as those harboured in deep sea hydrothermal vents, are able to survive, indeed flourish, under anoxic conditions through sulphur respirationanaerobic respiration with sulphur (10, 11). O2 levels gradually increased, and probably then fell, until around 700 million years ago when a further sharp increase occurred (12).

While O2 can be toxic in many respects, as discussed in a later section, its rise was critical to the development of multi-cellular organisms and the physiological complexity that this implies. The earliest known fossils of a eukaryote, from which multi-cellular organisms evolved, date from at least 2 billion years ago (14).

In eukaryotes and early multi-cellular organisms requiring O2, uptake occurs by direct transfer across the cell membrane in essentially the same manner as other tribology international journal prior to the development of specialised digestive and respiratory organs.

The ready availability of O2 had a profound effect on the metabolic opportunities for an organism and the consequent systems that developed. The mitochondrion, through respiration and oxidative phosphorylation, is a potent example of a cellular organelle whose evolution resulted in major new metabolic processes.

The most widely accepted view on the origin of mitochondria is the endosymbiotic hypothesis which proposes that mitochondria were originally prokaryotic cells (14, 15). These prokaryotes were able to undertake oxidative processes that early eukaryotic cells could not perform, and they subsequently became endosymbionts living within the eukaryote cell structure.

Animals are constant metabolisers, whether they are poikilotherms or homeotherms, and this is so even in those species that undergo periods of hibernation, aestivation or torpor (albeit at a reduced rate of metabolism). However, most nutrients are obtained in higher animals on an intermittent basis, such species being periodic feederswhether in i have to go now i have to go now form of distinct meals or through frequent foraging.

Foods, entering through the mouth, are generally complex structures and the nutrients that they contain are not immediately available. Instead, they require release through digestion and are subsequently absorbed from the gastrointestinal tract, a process that may involve specific transporters. In simple organisms, O2 is obtained in a manner similar to that of other nutrientsby absorption across the cell membranewhile in complex organisms it is fundamentally different. The evolution of specialised organs has resulted in the development of a respiratory system for the delivery of O2, differentiating it sharply from the route by which all other nutrients are provided through the i have to go now i have to go now system (Table 1).

This reflects both the constant metabolic need for O2 together with the absence of any significant storage. There is some limited storage, however, in skeletal muscle for local use through binding to the iron-containing protein myoglobin, but this is primarily a feature of marine animals such as whales, which experience apnoea intracerebral hemorrhage diving and where the haem protein is present in relative abundance (16). On entering the lungs, O2 passes into the alveoli which as highly vascularised sacs enable the rapid movement of the gas by simple diffusion, first across the alveolar epithelium and then the endothelial cells of the alveolar capillaries.

Once in the circulation, O2 binds to haemoglobin in the erythrocytes and is immediately transported to tissues (17). Modifications to this route of entry occur through the presence of gills in aquatic species, while in lower animals simpler systems for obtaining O2 are evident. The presence of haemoglobin as a specific carrier for O2 has some parallels with the transport and delivery of a number of other nutrients.

Once across the gastrointestinal wall, from mucosal to serosal side, nutrients move to their immediate sites of action or to storage organs hht subsequent use.

Storage occurs particularly in the liver and skeletal muscle for glucose as glycogen, and in white adipose tissue depots for the sequestration of fatty acids as triacylglycerols (19). In some cases, carrier proteins are involved in the transport of nutrients to their storage site, such as transferrin for the transport of iron to the bone marrow (21). Specific carriers, analogous to haemoglobin, also transport a number of nutrients to the tissues where they are required once released from storage, examples including retinol binding protein for retinol (21, 22) and plasma lipoproteins in the case of lipids (19, 23).

The central role of O2 as a nutrient is in mitochondrial respiration, acting as an electron acceptor thereby enabling ATP to be formed through oxidative phosphorylation. This process is fundamental to aerobic organisms, with the oxidation of glucose and fatty acids requiring the continuous provision of O2.

Several core metabolic pathways are Axitinib (Inlyta)- Multum to mitochondrial oxidative phosphorylationglycolysis, glycogenolysis, lipolysis, and the tricarboxylic acid (Krebs) cycle (19). White adipocytes, for example, have moderate numbers of mitochondria which contain limited cristae, with most of the volume of these cells being due to i have to go now i have to go now lipid droplet (25, 26).

Brown adipocytes, in marked contrast, contain large numbers of mitochondria with a highly developed and dense cristae structure, especially in rodents adapted to i have to go now i have to go now environments when maximum non-shivering thermogenesis state solid ionics required (25, 26).

In these circumstances, brown fat mitochondria utilise substantial amounts of O2 in order to sustain the oxidation of fatty acids and other substrates at high rates, with ATP synthesis being bypassed through a proton leakage pathway regulated by UCP1 (uncoupling protein-1) (27). The partial pressure of O2 is highest at sea level, but falls with altitude leading to a decrease in the amount available.

Altitude is one of the several environmental situations that result in a reduction in the availability of O2. Animals, including humans, that habitually live at high elevations have evolved distinct physiological adaptations which allow them to adapt to the relatively hypoxic conditions.

Another environmental circumstance in which O2 deprivation occurs, albeit on a short-term basis, is that sexual orientation quiz by aquatic mammals such as whales during deep sea dives.



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