Ectothermy and endothermy

All animals thermoregulate. The internal environment of the body is under the influence of both external and internal conditions. Land animals thermoregulate in several ways. They do so behaviorally, by moving to a colder or warmer place, by exercising to generate body heat, or by panting or sweating to lose it. They also thermoregulate physiologically, by activating internal metabolic processes that warm or cool the blood. But these efforts have limits, and, as a result, external temperatures and climatic conditions are among the most important factors controlling the geographic distribution of animals.

Today’s so-called warm-blooded animals are the mammals and birds; reptiles, amphibians, and most fishes are called cold-blooded. These two terms, however, are imprecise and misleading. Some “cold-blooded” lizards have higher normal body temperatures than do some mammals, for instance. Another pair of terms, ectothermy and endothermy, describes whether most of an animal’s heat is absorbed from the environment (“ecto-”) or generated by internal processes (“endo-”). A third pair of terms, poikilothermy and homeothermy, describes whether the body temperature tends to vary with that of the immediate environment or remains relatively constant.

Today’s mammals and birds have a high metabolism and are considered endotherms, which produce body heat internally. They possess biological temperature sensors that control heat production and switch on heat-loss mechanisms such as perspiration. Today’s reptiles and amphibians, on the other hand, are ectotherms that mostly gain heat energy from sunlight, a heated rock surface, or some other external source. The endothermic state is effective but metabolically expensive, as the body must produce heat continuously, which requires correspondingly high quantities of fuel in the form of food. On the other hand, endotherms can be more active and survive lower external temperatures. Ectotherms do not require as much fuel, but most cannot deal as well with cold surroundings.

From the time of the earliest discoveries in the 19th century, dinosaur remains were classified as reptilian because their anatomic features are typical of living reptiles such as turtles, crocodiles, and lizards. Because dinosaurs all have lower jaws constructed of several bones, a reptilian jaw joint, and a number of other nonmammalian, nonbirdlike characteristics, it was assumed that living dinosaurs were similar to living reptiles—scaly, cold-blooded, ectothermic egg layers (predominantly), not furry, warm-blooded live-bearers. A chauvinistic attitude seems to prevail that the warm-bloodedness of mammals is better than the cold-blooded reptilian state, even though turtles, snakes, and other reptiles do very well regulating their body temperature in a different way. Moreover, both birds and mammals evolved from ectothermic, poikilothermic ancestors. At what point did metabolism heat up?

Clues to dinosaurian metabolism

The question of whether any extinct dinosaur was a true endotherm or homeotherm cannot be answered, but some interesting anatomic facts suggest these “warmer” possibilities. Probably the most direct evidence of dinosaurian physiology comes from bones themselves, particularly in regard to how they grew. The long bones (such as arm and leg bones) of most dinosaurs are composed almost exclusively of a well-vascularized type of bone matrix (fibro-lamellar) also found in most mammals and large birds. This type of bone tissue always indicates rapid growth, and it is very different from the more compact, poorly vascularized, parallel-fibred bone found in crocodiles and other reptiles and amphibians. It is generally thought that well-vascularized, rapidly growing bone can be sustained only by high metabolic rates that bring a continual source of nutrients and minerals to the growing tissues. It is difficult to explain these histological features in any other metabolic terms. On the other hand, most dinosaurs retain lines of arrested growth (LAGs) in most of their long bones. LAGs are found in other reptiles, amphibians, and fishes, and they often reflect a seasonal period during which metabolism slows, usually because of environmental stresses. This slowdown produces “rest lines” as LAGs in the bones. The presence of these lines in dinosaur bones has been taken as an indication that they were metabolically incapable of growing throughout the year. However, LAGs in dinosaurs are less pronounced than in other reptiles; LAGs can also appear in different numbers in different bones of the same skeleton, and they are sometimes even completely absent. Finally, some living birds and mammals, which are clearly endotherms, have LAGs very much like those of dinosaurs, so LAGs are probably not strong indicators of metabolism in any of these animals.

Other, less direct lines of evidence may reveal other clues about dinosaurian metabolism. Two dinosaurian groups, the hadrosaurs and the ceratopsians, had highly specialized sets of teeth that were obviously effective at processing food. Both groups were herbivorous, but unlike living reptiles they chopped and ground foliage thoroughly. Such highly efficient dentitions may suggest a highly effective digestive process that would allow more energy to be extracted from the food. This feature by itself, however, may not be crucial. Pandas, for example, are not very efficient in digesting plant material, but they survive quite well on a diet of almost nothing but bamboo.

Another line of evidence is that dinosaurs had anatomic features reflecting a high capacity for activity. The first dinosaurs walked upright, holding their legs under their bodies; they could not sprawl. This indicates that, by standing and walking all day, they probably expended more energy than reptiles, which typically sit and wait for prey. As some lineages of dinosaurs grew larger, they reverted to four-legged (quadrupedal) locomotion, but their stance was still upright. They also put one foot directly in front of the other when they walked (parasagittal gait), instead of swinging the limbs to the side. Such posture and gait are present in all nonaquatic endotherms (mammals and birds) today, whereas a sprawling or semierect posture is typical of all ectotherms (reptiles and amphibians). Bipedal stance and parasagittal gait are not sustained in any living ectotherm, perhaps because they require a relatively higher level of sustained energy.

The high speeds at which some dinosaurs must have traveled have also been invoked as evidence of high metabolic levels. For example, the ostrichlike dinosaurs, such as Struthiomimus, Ornithomimus, Gallimimus, and Dromiceiomimus, had long hind legs and must have been very fleet. The dromaeosaurs, such as Deinonychus, Velociraptor, and Dromaeosaurus, also were obligatory bipeds. They killed prey with talons on their feet, and one can argue that it must have taken a high level of metabolism to generate the degree of activity and agility required of such a skill. However, most ectotherms can move very rapidly in bursts of activity such as running and fighting, so this feature may not provide conclusive evidence either.

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Related to the upright posture of many dinosaurs is the fact that the head was often positioned well above the level of the heart. In some sauropods (Apatosaurus, Diplodocus, Brachiosaurus, and Barosaurus, for instance), the brain must have been several metres above the heart. The physiological importance of this is that a four-chambered heart would be required for pumping freshly oxygenated blood to the brain. Brain death follows very quickly when nerve cells are deprived of oxygen, and to prevent it most dinosaurs must have required two ventricles. In a four-chambered heart, one ventricle pumps oxygen-poor venous blood at low pressure to the lungs to absorb fresh oxygen (high pressure would rupture capillaries of the lungs). A powerful second ventricle pumps freshly oxygenated blood to all other parts of the body at high pressure. To overcome the weight of the column of blood that must be moved from the heart to the elevated brain, high pressure is certainly needed. In short, like birds and mammals, many dinosaurs apparently had the required four-chambered heart necessary for an animal with a high metabolism.

The significance of thermoregulation can be seen by comparing today’s reptiles with mammals. The rate of metabolism is usually measured in terms of oxygen consumed per unit of body weight per unit of time. The resting metabolic rate for most mammals is about 10 times that of modern reptiles, and the range of metabolic rates among living mammals is about double that seen among reptiles. These differences mean that endothermic mammals have much more endurance than their cold-blooded counterparts. Some dinosaurs may have been so endowed, and although they seem to have possessed the cardiovascular system necessary for endothermy, that capacity does not conclusively prove that they were endothermic. There exists the possibility that dinosaurs were neither complete ectotherms nor complete endotherms. Rather, they may have evolved a range of metabolic strategies, much as mammals have (as is illustrated by the differences between sloths and cheetahs, bats and whales, for example).