It is well known that the productivity of an animal is not necessarily identical with its fecundity. Fish or frogs which lay thousands of eggs at a time (a pike, for example, produces 130,000) are obviously more prolific than, for instance, a sparrow which lays only 18 eggs in a year, or than a rabbit, which in the same time gives birth to from 25 to 50. However, to produce this much smaller quantity of eggs or of young, the sparrow and the rabbit (I have chosen the most prolific bird and mammal) expend a much larger quantity of material than the frog or the fish. The sparrow and the rabbit employ in producing their progeny a bulk of material greater than the weight of their body, whilst the enormous quantity of eggs laid by the frog does not weigh more than one-seventh part of the body of the frog. It may be laid down, as a general rule, that although fecundity, that is to say the number of eggs or of young which are produced, diminishes as the organism becomes more complex, the productivity on the other hand increases, expressed in percentage of weight. The productivity, which is not more than 18 per cent. in batrachia, reaches 50 per cent. in reptiles, 74 per cent. in mammals, and 82 per cent. in birds.
It is plain that if reproduction shortens the life of mammals by weakening the organism, it must be the productivity, not the fecundity, which is the important factor. I have just shown that productivity is greater in birds than in mammals, and in consequence it cannot be on account of any greater burden of reproduction that mammals have a shorter life than birds. The shortness of mammalian life, again, cannot be attributed to the fact that mammals give birth to young, whilst the long-lived reptiles and birds produce eggs, because the longevity of the males, which produce neither young nor eggs, is none the less practically equal to that of the females of the same species. The reason of the short life of mammals must be sought for elsewhere.
III
THE DIGESTIVE SYSTEM AND SENILITY
Relations between longevity and the structure of the digestive system—The Cæca in birds—The large intestine of mammals—Function of the large intestine—The intestinal microbes and their agency in producing auto-intoxication and auto-infection in the organism—Passage of microbes through the intestinal wall
We have seen that the duration of life in mammals is relatively shorter than that in birds, and in the so-called “cold-blooded” vertebrates. No indication as to the cause of this difference can be found in the structure of the organs of circulation, respiration, or urinary secretion, or in the nervous or sexual apparatus. The key to the problem is to be found in the organs of digestion.
In reviewing the anatomical structure of the digestive apparatus in the vertebrate series, one soon comes to the striking fact that mammals are the only group in which the large intestine is much developed. In fish, the large intestine is the least important part of the digestive tube, being little wider in calibre than the small intestine. Amongst batrachia, where it is a relatively wide sack, it has begun to assume some importance. In several reptiles it is still larger, and may be provided with a lateral out-growth, which is to be regarded as a cæcum. In birds, the large intestine still remains relatively badly developed; it is short and straight. In most birds, at the point where the large intestine passes into the small intestine, there is a pair of cæca, more or less developed. These cæca are absent in climbing birds, such as the wood-pecker, the oriole, and many others. They are reduced to a pair of tiny outgrowths in the eagles, sparrow-hawks, and other diurnal birds of prey, and in pigeons, and perching birds. These organs are larger in the nocturnal birds of prey, in gallinaceous birds, and in ducks, etc.[40]
In the large running birds, such as ostriches, rheas, and tinamous, the cæca are relatively largest. Thus, for instance, in a rhea (Rhea americana) which I dissected, the cæca were nearly two-thirds as long as the small intestine. The latter was 1·65 m. in length, whereas one of the cæca was 1·01 m., and the other 0·95 m. The weight of the two cæca with their contents was more than 10 per cent. of the total weight of the bird.
Notwithstanding the exceptions, which are relatively rare, the large intestine is badly developed in the case of birds. On the other hand, it reaches its largest size amongst mammals. In these animals, “only the posterior portion of the latter, or rectum, which passes into the pelvic cavity, corresponds to the large intestine of lower Vertebrates; the remaining, and far larger part, must be looked upon as a neomorph, and is called the colon.”[41]