Friday, 22 February 2013

Breathing and warm-bloodedness – another old post

Having just started on my science post, and realising it will be longer than expected, I figure I should feed you some of the last useful posts from my old website. Also, this Monday we are having a one-day fieldtrip (we just found out...), so I might bring something worth telling from there as well. Meanwhile, I apologise for the time it is taking to write the big thing up, but stuff keeps coming up that need to be done – such is university

Tetrapods (land-living vertebrates) breathe because our internal chemical reactions consume oxygen. Without oxygen, we would simply not be able to live. (This should not surprise you.) 

The chemical reactions in question are described in detail in the article about aerobic and anaerobic metabolism. Here, I will briefly summarise the key points. Aerobic metabolism refers to a series of chemical reactions involved in aerobic cell respiration, which basically uses oxygen to release energy from food for the cells to carry out their functions. There is an alternative way of extracting energy from food without the use of oxygen – anaerobic cell respiration – but it consumes about twenty times as much food to provide a given amount of energy. 

There is a limit to how much oxygen an animal can absorb; this depends on its lung capacity. A large influx of oxygen is essential, because when the oxygen supply cannot meet the energy demand of the cells (e.g. during intense activity), anaerobic cell respiration must be used to supply the additional energy required – which is undesirable since it consumes much more food. 

Lung capacity can partly be related to posture. Dinosaurs, just like mammals and birds (as well as some archosaurs, such as pterosaurs and rauisuchians) had an erect or parasagittal limb posture – they held their limbs vertically under their bodies (seen from the front). This is contrasted by the sprawling limb arrangement of most other reptiles, and amphibians, where the upper bones of the arms and legs are held almost horizontally and pointing away from the trunk.





Animals with a sprawling posture move very differently from those with a parasagittal limb arrangement. Sprawlers move by alternately contracting their left and right flanks. This helps moving the limbs forward, making each step considerably longer, but at a fatal cost: when contracting each side, the lungs are also compressed, preventing air from flowing in. Consequently, sprawlers cannot breathe while they run. Thus, their lung capacity, and therefore also oxygen influx, is severely limited during strenuous activity – precisely when they need oxygen the most. When animals with parasagittal limbs move, the lung capacity is not affected. There is, of course, still a limit to how much oxygen they can extract, but the lungs are not impeded by movement, allowing them to maintain a continuous inflow of oxygen, reducing the need for anaerobic supplement. 

How does this relate to warm-bloodedness, then? The above example illustrates that animals with a parasagittal limb posture can rely on a predominantly aerobic metabolism – i.e. they extract their energy mostly by aerobic cell respiration. This allows them to extract more energy from a given amount of food. 

Modern animals with parasagittal limbs (mammals and birds) exploit this advantage by spending a substantial amount of energy on producing heat internally. They can afford this because their energy economy is so efficient. The heat is produced by speeding up chemical reactions in their cells – they are said to have a high basal metabolic rate (BMR), or being tachymetabolic. In other words, they are actively consuming considerable amounts of food even when at rest, in order to keep their bodies warm. Their efficient respiratory system enables them to maintain high metabolic rates even when engaged in active movement.  Sprawlers, however, cannot have high BMRs because their respiratory system is too limited (either, they are not able to maintain elevated metabolic rates even at rest, or the additional demand during strenuous action simply becomes too much). 

Another way the parasagittal limb posture is linked to warm-bloodedness is by activity levels. Warm-blooded animals have a superb advantage over cold-blooded ones because they can be active for a considerably longer proportion of the day. Cold-bloods tend to spend a lot of time lying still, either in ambush or for behavioural thermoregulation (sun basking to warm up, or cooling down in the shade), while mammals and birds are active for a substantial proportion of the day (or night). And a parasagittal stance is designed for being capable of long-term, energy-efficient activity.  

There is nothing saying that dinosaurs must have been tachymetabolic and/or highly active because they had a parasagittal limb posture. But the fact that the only tachymetabolic animals today are those with parasagittal limb arrangement suggests that there is a tight relationship. We may infer that dinosaur at least were capable of being warm-blooded thanks to their erect stance. 

Birds have the by far most efficient respiratory (breathing) system known. For a detailed account of this system, please see the article on birds. In brief, they rely on so-called air sacs – cavities in their bones – where they pass the air after gas exchange (i.e. oxygen has been replaced with carbon dioxide), before it is exhaled; this allows the birds to maintain a continuous flow of air through the lungs – in contrast to other tetrapods, where the lungs are alternately filled and emptied – and so enables optimal oxygen extraction from the air. It is thanks to this ingenious design that birds can fly across oceans in their long seasonal migrations: not only does it allow for extensive aerobic respiration; it also makes their skeletons lighter. 

Ample fossils of dinosaurs with pneumatised bone have been found. This bone contains air cavities like those of birds, inviting the idea that dinosaurs possessed a similar, if not identical respiratory system to that of modern birds. Although few dinosaurs flew, impressive stamina and a light body can provide many advantages, such as allowing predators to engage in fast, lengthy pursuit of prey. 

However, since mammals do not possess such adaptations, bone pneumatisation does not have a strong connection with warm-bloodedness. It is not a requirement, but more of an extension to enhance the efficiency of the fundamental feature – aerobic metabolism. Pneumatic bones supports the idea of dinosaurs as using aerobic metabolism to a considerable extent – perhaps more, perhaps less, than modern warm-bloods. On the other hand, this is not compulsory: it is possible that the sole purpose of the pneumatisation was weight reduction, which may have been important for the larger dinosaur forms. However, this line of reasoning does not explain why the bone pneumatisation was most well-developed in the smaller forms! 

A final piece of evidence related to the aspect of breathing is the possession of a secondary palate – the structure dividing the nasal and oral cavities and so enables breathing while chewing – by hadrosaurids and ankylosaurids, as well as mammals. This is highly useful for herbivores employing extensive chewing of their food. Without the secondary palate, there would have been a greater risk of food ending up in the lungs, if the animal would try to breathe during mastication. The ability to chew and breathe at the same time removes the need to compromise between food and oxygen intake, both being crucial to sustaining a high metabolic rate. 

Now, a secondary palate is by no means necessary for warm-blooded herbivores: birds do not possess them. Moreover, cold-blooded animals such as crocodiles and turtles also have secondary palates, making arguments for dinosaurian warm-bloodedness based on this correlation very weak. Rather, it seems that the secondary palate, like bone pneumatisation, would improve warm-bloodedness, if already present. (Note that it can improve the conditions of cold-blooded animals too – recall the crocodiles and turtles!)

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