Breath-holding in non-human animals

Lin (1982) reported on bradycardia studies with dogs, using dogs which showed an ability to hold their breath. ("Breath-hold Diving in Terrestrial Mammals" by Yu-Chong Lin (Department of Physiology, John A. Burns School of Medicine, University of Hawaii, Honolulu, Hawaii). In Exercise and Sport Sciences Review, 1982, vol. 10, pp. 270-307.)

By the way, seals and whales don't hold their breath when they dive; they store oxygen in their blood, and actually expel the air from their lungs as they dive. This system -- radically different from humans -- is used by pinnipeds and cetaceans.

The Descended Larynx and the AAT/H

The descended larynx of humans therefore:
A) undergoes this change during precisely the period when infants learn to speak -- in an ever more complex manner; and
B) isn't fully formed for the first four years of the infant's life.

If it were an adaptation which "aquatic" hominids had developed from living in their environment, these "aquatic" infants would without its supposed benefits for the first four years of their lives. And of course, since one of the unfortunate consequences of this feature is that we can get water into our lungs more easily, it's difficult to see why a feature which promotes drowning would be considered "aquatic".

The idea that this is an aquatic feature also doesn't explain why, in human males, the larynx descends even further at puberty (that's why male voices get dramatically lower at that time). A far more sensible reason, based on research rather than speculation, will show up in a few paragraphs.

Even some AAT/H proponents, such as Marc Verhaegen, have realized that this feature is not at all like the larynx of the vast majority of aquatic mammals, although he still has described it as an "aquatic" trait. When Elaine Morgan acknowledged Verhaegen's finding this out, she also admitted she just hadn't looked at the evidence on this -- excuse me? didn't look at the evidence? (that isn't someone I'd want to buy a used theory from...)

Morgan has stated that the descended larynx indicates habitual mouth breathing, which she says, without evidence, would be better for diving, but we see that other terrestrial mammals mouth breath just as humans do when exerting themselves -- common mammals like dogs and cats, so that doesn't add up. Other than that, it's merely suggested that it must be an aquatic trait because it's seen only in humans and (a very few) aquatic mammals. In short, there is no good explanation given why this would be an aquatic trait, and it turns out that there's excellent reasons to see that it isn't, for instance, the fact that, contrary to common AAT/H claims, there are other terrestrial mammals with descended larynxes.

Previously on this page I had mentioned a problem concerning the descended larynx as a feature of our transitional population (as Morgan claims it) -- that the skulls of hominids after the transition don't show the basicranial angle we see in humans that indicates a fully descended larynx. It occurred several million years later. However, it's possible that a descended larynx wouldn't leave this mark in the basicranial angle, but even this does nothing to comfort the AAT/H proponent who uses the larynx as an example of an aquatic trait, because it turns out that there are several terrestrial mammals (and even some birds) with permanently descended larynxes, and even more with larynxes that habitually descend when vocalizing.

For instance, chimpanzees have been found to have partially descended larynxes -- that's right, our closest relatives are similar to us in this feature, although their hyoid (the U-shaped bone that supports the tongue muscles) doesn't also descend as ours does. (This gives us more fine control of vocalization than they have.)

Nishimura, Mikami, Suzuki, and Matsuzawa explain in the abstract of their 2003 paper ("Descent of the larynx in chimpanzee infants"):

We used magnetic resonance imaging to study the development of three living chimpanzees and found that their larynges also descend during infancy, as in human infants. This descent was completed primarily through the rapid descent of the laryngeal skeleton relative to the hyoid, but it was not accompanied by the descent of the hyoid itself.

Comparative Vocal Production and the Evolution of Speech: Reinterpreting the Descent of the Larynx (2002)
W. Tecumseh Fitch:

9th pg.: The work on animal vocal anatomy dates mostly from the 19th century, and was based exclusively upon dissections of dead animals. By the time new techniques allowing anatomical visualization of living animals were developed, such as x-ray film (cineradiography) or MRI, the study of comparative anatomy had fallen from favor. Although occasional critics pointed out that the crucial issue is not the resting position of the larynx, but its position during vocalization (Nottebohm 1976), it is only recently that modern imaging techniques have been applied to live vocalizing mammals (Fitch 2000a). These data indicate that mammals lower the larynx as a matter of course during vocalization, in some cases approaching the "two-tube" configuration typical of adult humans. Furthermore, new anatomical analyses show that humans are not unique in our laryngeal position, because several other species also have a permanently descended larynx.
Comparative Vocal Production and the Evolution of Speech: Reinterpreting the Descent of the Larynx W. Tecumseh Fitch

9th pg.: These data indicate that the vocal tract configuration of vocalizing animals, at least in dogs, pigs, goats and monkeys, is more similar to that of human talkers than was previously inferred on the basis of dissections of dead animals. In particular, all four nonhuman species examined can and do lower their larynges into the oral cavity during loud vocalizations, either to a relatively minor degree (goats) or to a surprisingly extensive degree (dogs).
W. Tecumseh Fitch

11th pg.: Finally, these data suggest that a lowered larynx during vocalization is in fact a primitive trait that we share with other mammals, rather than a uniquely human adaptation. What is unusual about our species is that the adult human larynx is permanently lowered, rather than dropping only during vocalization. However, other recent data show that even this difference is not uniquely human.
W. Tecumseh Fitch

11th pg.: But despite the claims of Negus (1949), this position is not unique to humans. In at least two species of deer, red deer Cervus elaphus and fallow deer Dama dama, the larynx of postpubertal males is permanently lowered to a resting position comparable to that in humans.
W. Tecumseh Fitch

How about the diving reflex: are humans well adapted for diving?

AAT/H proponents say they do, and that other terrestrial mammals don't show such responses as breath-holding and the "diving" reflex. The following scientists' writings show that the facts say otherwise. Note also that although Hardy said in 1960 that the diving reflex was only found in humans and diving animals, contrary evidence had been available for over 20 years before that.

1982 "Breath-hold Diving in Terrestrial Mammals" by Yu-Chong Lin (Department of Physiology, John A. Burns School of Medicine, University of Hawaii, Honolulu, Hawaii). In Exercise and Sport Sciences Review, vol. 10, pp. 270-307.
 

pg. 271: However, universality of the "diving" responses is well recognized in that these responses exist in all vertebrates studied so far, differing only in intensity and tempo.

pg. 272: Many previous reports indicated that there is a slow development of bradycardia response in humans. Undoubtedly, these reports refer not to the onset but to the slow attainment of the maximum bradycardia response. It should be noted that diving mammals arrive at maximum bradycardia much faster.

pg. 277: In contrast to diving species, hypertension is a consistent finding in man during BH.

1983 Diving and Asphyxia: A comparative study of animals and man by Robert Elsner (Professor of Marine Science, University of Alaska) and Brett Gooden (Adelaide, South Australia. Formerly Lecturer in Physiology, University of Nottingham). Cambridge University Press: Cambridge.
 

pg. 20: Bradycardia during diving, breath-holding and asphyxia has been observed in many vertebrate species, including man. Among those studied are rabbit (Bauer, 1938), sloth (Irving, Scholander & Grinnell, 1942b), snake (Johansen, 1959; Murdaugh & Jackson, 1962), dog (Elsner et al., 1966a), pig (Irving, Peyton & Monson, 1956), armadillo (Scholander, Irving, Peyton & Grinnell, 1943), echidna (Augee et al., 1971) and lizards (Wood & Johansen, 1974). Terrestrial mammals generally do not respond as immediately nor as markedly as the aquatic species.

1990 Diving Medicine, Second Edition. by Alfred A. Bove, M.D. (Chief, Section of Cardiology, Temple University Hospital, Philadelphia, Pennsylvania) and Jefferson C. Davis, M.D. (Hyperbaric Medicine, P.A., San Antonio, Texas). W.B. Saunders Company: Philadelphia, London, Toronto, Montreal, Sydney, Tokyo.

pg. 66: Unlike in diving animals, arterial blood pressure seems to increase while cardiac output decreases only slightly during breath-hold diving in humans.

pg. 66: One of the most intriguing findings with regard to diving medicine is the occurrence of cardiac arrhythmias during breath-hold diving. This is in contrast to diving animals who display a marked bradycardia but never show any cardiac arrhythmias during diving. In humans, it is common to find various types of arrhythmias even during simply breath-holding.

1992 Diving and Subaquatic Medicine, Third Edition. by Carl Edmonds, Christopher Lowry, and John Pennefather.

pg. 357: In humans, the diving reflex is more rudimentary and undeveloped. Although there is a diving bradycardia, it is often complicated by the development of idioventricular foci producing ectopic beats. ECG abnormalities are frequent during or after the dive. T-wave inversion, premature ventricular excitation and atrial fibrillation, together with other irregularities and dysrhythmias, are common.

pg. 358: The drop in cardiac output noted in the diving mammals is seen to only a slight degree in humans.

1990 Sea Otters by Marianne Riedmann. Monterey Bay Aquarium: Monterey, California

pg. 33: While diving for food, sea otters must conserve oxygen and contend with underwater pressure. They generally forage in shallow waters of less than 60 feet, and their feeding dives usually last one to two minutes. Yet sea otters have been known to dive to depths of 330 feet and remain under water for up to four or five minutes. In contrast, a person without oxygen passes out within three minutes. Sea otters can hold their breath for such a long time in part because their lungs are nearly two-and-a-half times larger than the lungs of a similar-sized mammal. Large lungs help store oxygen and regulate buoyancy while the otter floats on the surface. And sea otters have flexible ribs that allow the lungs to collapse under pressure. Otters also have cartilaginous airways connected directly to tiny, air-filled lung sacs which help provide an unrestricted flow of oxygen to the blood. In addition, the sea otter's blood has a higher buffering capacity than that of non-diving mammals, which helps the otter handle the excess carbon dioxide that accumulates under pressure during a dive.

pp. 21-22: Like seals, sea otters have a higher concentration of haemoglobin than terrestrial mammals (and river otters) which enables them to carry more oxygen in their blood. They also have relatively large lungs which, compared to other marine mammals, are twice as large as expected and are the reason for the sea otter's large rib cage. The lungs may form an important oxygen store but it has been suggested that their importance is in providing buoyancy rather than an air supply. This is very necessary in an animal which has little or no fat to give buoyancy and which often carries large stones with it when feeding and foraging (see Chapter 3).

1990 The Pinnipeds: Seals, Sea Lions, and Walruses by Marianne Riedmann. University of California Press: Berkeley, L.A., and Oxford.

pp. 28-29: Bradycardia and the channeling of blood to critical organs save oxygen. Since muscles are deprived of their usual supply of blood and oxygen, they compensate by containing large amounts of a special compound called myoglobin, an iron-bonding pigment related to hemoglobin that stores oxygen. Myoglobin helps the seal to tolerate the large accumulation of carbon dioxide in its bloodstream that prompts breathing during periods of prolonged breath-holding. Myoglobin therefore helps the animal to conserve oxygen during high-speed movement or deep dives. In fact, cetacean and pinniped muscles appear nearly black when they are exposed to the air. This dark coloration is due to the extremely dense concentrations of deep red myoglobin in the muscle tissues.

A seal's muscles are also able to handle the high amounts of lactic acid that accumulates in its system during periods of heavy or prolonged activity (such as diving), which causes muscle exhaustion in land mammals. The muscles of marine mammals can function with insufficient oxygen for many hours. Most of the time, the muscles function aerobically (with oxygen), even though the seal may be diving and exercising heavily. They function anaerobically when oxygen supplies are depleted by prolonged submergence. When a seal surfaces from a long dive, it takes several rapid and deep breaths until the body's oxygen supply is replenished; at the same time, at least in harbor seals, a pronounced tachycardia (extremely rapid heartbeat) occurs (Fedak et al. 1985). Seals and whales even have an automatic mechanism that cuts off breathing if they are knocked unconscious, so that their lungs will not fill with water.

In pinnipeds, the oxygen-transporting circulatory system is very large. A seal total blood volume in relation to its body weight is 1.5-2 times greater than that of other mammals. According to Scheffer (1976), "In a full-grown walrus the forked veins which drain the lower body are so enormous that a man can draw them over his legs like a pair of pants!" Wickham et al. (1985) and others (e.g., Lenfant et al. 1970) have identified a number of characteristics of phocid seal blood that appear to help the seal to cope with the hypoxia, or oxygen deficiency, of long dives. Phocid blood has a greater capacity than the blood of terrestrial mammals to store oxygen because phocids have fewer and larger red blood cells with a higher concentration of hemoglobin, which stores and carries oxygen. And despite the high viscosity, or thickness, of phocid blood, the rate of blood flow is high.

1990 The Natural History of Seals by W. Nigel Bonner. Facts on File: New York, Oxford, and Sydney.
 

pp. 33-34: Breath-hold capacity can be increased by taking more oxygen down with each dive. A human diver will breathe deeply -- hyperventilate -- before diving, and descend with lungs full of air. There are disadvantages in this. For a start, full lungs increase buoyancy and so make it more difficult to descend; secondly, there are pressure problems associated with diving with full lungs. Seals hyperventilate before diving, but they expel most of the air from their lungs before they go down, and hence they do not depend on air from the lungs for their oxygen when they dive.

Besides being stored as a gas in the lungs, oxygen can be carried in physical solution in the blood and tissue fluids, or chemically bound to haemoglobin in the red blood cells or to another respiratory pigment, myoglobin, in the muscles. In seals the major store is carried in the blood. Seals have greater blood volumes than terrestrial mammals. A Weddell seal, for example, has about 150 ml of blood per kilogram bodyweight, about twice the value for Man. Furthermore, seal blood contains more haemoglobin than human blood -- about 1.6 times as much. The combined result of this is that the blood oxygen stores per unit of body weight is about three or three and a half times that of Man. Gerry Kooyman (1981) has pointed out that because blubber is more or less inert and represents about 30 percent of the seal's body weight, a more realistic comparison might be to lean body weight, in which case a seal's blood oxygen store would be about 5.3 times that of Man (though it should be borne in mind that an obese human may have about the same proportion of fat -- though differently distributed -- as a seal). Not only does the seal have more haemoglobin in its blood than Man, it also has very much more myoglobin in its muscles. This protein, which resembles haemoglobin in its ability to combine loosely with oxygen, is what gives muscles their red color.  [my notes here unfortunately left out a partial sentence. sorry; the following statement refers to the muscles of seals] Weddell, containing about ten times the myoglobin concentration of human muscle, is almost black.