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Death Squared: the Explosive Growth and Demise of a Mouse Population

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Death Squared: the Explosive Growth and Demise of a Mouse Population 80 Proc. roy. Soc. Med. Volume 66January 1973 peculiar to man allows for the learning of Death Squared: language and social habits, and is undoubtedly the result of selection of factors making for The Explosive Growth and Demise delayed adolescence, enabling the more co- of a Mouse Population operative groups ofpeople to survive. by John B Calhoun MD It is an interesting question whether still closer (Section on Behavioral Systems, social co-operation might be achieved by a society Laboratory ofBrain Evolution & Behavior, that adopted some method of clonal reproduction National Institute ofMental Health, by which all men were truly brothers. Altruism is 9000 Rockville Pike, an effective evolutionary policy only if it benefits Bethesda, Maryland20014, USA) your own genes, and the ants and bees use very special methods to ensure this. Technically such I shall largely speak of mice, but my thoughts are changes begin to look almost possible; whether on man, on healing, on life and its evolution. they are desirable is another matter. We certainly Threatening life and evolution are the two deaths, cannot foresee them in the near future, but the death of the spirit and death of the body. Evolu- possibility of them may remind us that the tion, in terms ofancient wisdom, is the acquisition biggest difference of all between men and animals of access to the tree of life. This takes us back to is in the vast range of techniques of control of the white first horse of the Apocalypse which with life that are now becoming available. We are all its rider set out to conquer the forces that familiar with the effects of the techniques of threaten the spirit with death. Further in Revela- physical science in promoting the industrial tion (ii.7) we note: 'To him who conquers I will revolution and the invention ofmachines. Perhaps grant to eat the tree' of life, which is in the it is still insufficiently realized that biological paradise of God' and further on (Rev. xxii.2): knowledge is now producing a still greater revolu- 'The leaves of the tree were for the healing of tion, namely in the way we manage ourselves. nations.' Inevitably the medical profession finds itself in This takes us to the fourth horse of the the forefront of this revolution and has to learn Apocalypse (Rev. vi.7): 'I saw ... a pale horse, to deal with all the new problems that it brings. and its rider's name was Death, and Hades This symposium is especially welcome for that followed him; and they were given power over a reason. I should like to emphasize once more that fourth of the earth, to kill with the sword and with the problems are those of information and con- famine and with pestilence and by wild beasts of trol. We shall collect much information today. the earth' (italics mine). This second death has We shall probably not have much to say about gradually become the predominant concem of the problems ofhow it is to be used for the control modern medicine. And yet there is nothing in the of human life. But I hope that we shall not forget earlier history of medicine, or in the precepts that if we are to achieve successful regulation in embodied in the Hippocratic Oath, that precludes the future we shall have to be ready to use and medicine from being equally concerned with to accept control in matters of population and healing the spirit, and healing nations, as with environment. The need for such control should healing the body. Perhaps we might do well to not alarm us, ifit is properly based on information reflect upon another of John's transcriptions and feed-back. After all, it is a natural extension (Rev. ii. 1): 'He who conquers shall not be hurt of the regulation by the information of our DNA by the second death.' or by the language and social customs that we inherit from the past. Bodily Mortality We should never forget that the continuation Let us first consider the second death (Table 1). of our lives depends every moment on information The four mortality factors listed in Revelation from the past. We are as we are because of the have direct counterparts (with a division of one endowment we receive, particularly from our of them to form a total of five) in the ecology of genes, our language and our social system. With- animals in nature. I shall briefly treat each of out them we are nothing. Life depends upon regulation, and the opposite of control is death. Table 1 We should be thankful that we are subject to The second death genetic, linguistic and social systems that allow us however far it be As in Revelation vi.8 Ecological expression a full and developing life, may (1) Sword (1) Emigration from perfection, whatever that may mean. (2) Famine (2a) Resource shortage Let us rejoice in the fact that we live at a time (2b) Inclement weather (and fire and cataclysms when ever-fresh sources of information abound, ofnature) and when people are beginning to realize the need (3) Pestilence (3) Disease for closer and closer co-operation for survival. (4) Wild beasts (4) Predation Symposium on Man in HisPlace 81 Fig 1 Universe 25, February 101970, 681 days after colonization. Calhoun in universe with an exceptionally large aggregate ofpooled withdrawn mice on thefloorjust across the centre ofthefloorfrom him. Note the aggregation ofmice on somefood hoppers in contrast to their near absence on others. This is the 'behavioural sink' phenomenon in which the learnedneedfor proximity to others as a secondary reinforcer at a resource site gains dominance over theprimary need, in this casefood. Photo by Yoichi R Okamoto these five mortality factors, and then discuss the (2b) Inclement weather: Every species of animal steps taken to eliminate, or drastically reduce, the has developed a genetic adaptation to some impact of each in a Utopian environment con- particular range of external conditions which structed for mice. affect its physiology. Any conditions of wind, rain, humidity or temperature which exceed the (1) Emigration: Animals in the wild rarely die usual limits of tolerance bring death immediately from the 'sword' directly; few are the deaths from or increase the risk of death through debilitation. intraspecific combat. Rather it is that individuals, Beyond these more usual contributions to mor- who have failed in the more symbolic conflicts tality, flood and fire represent cataclysmic changes involved in gaining the right to remain in the that have widespread and more long-lasting effects locality of their birth or in more preferred on population numbers. habitats, take recourse to movement into peri- (3) Disease: Although most animals develop some pheral unoccupied or suboptimal habitats. In capacity to tolerate the parasites, bacteria and strange and less favourable habitats the emigres viruses that invade their bodies, most species still become more exposed to other mortality factors. remain subject to epidemic decimation in addition Any removal of excess members from an estab- to a more normal attrition from disease. Abnor- lished population, from the point of view of mally high densities enhance the likelihood of change in its numbers, represents mortality. In spread of disease to epidemic proportions. this sense emigration is a mortality factor. (4) Predation: Practically every species has, (2a) Resource shortage: Classically food and water through evolution, had associated with it preda- have received the focus of attention with respect tors capable ofkilling some ofits members. to their reduction increasing the likelihood of death. Shortages of shelter, other environmental Despite loss of members from these five kinds resources and associates lead to debilitation, and of mortality, most species persist over long an unsatisfactoriness of habitat that culminate in periods, even those measured in geological time. death or failure to reproduce (species death). To do so every such species has developed 82 Proc. roy. Soc. Med. Volume 66January 1973 capacities for reproduction requisite to compen- until the population exceeded 3840. Due to the sate for losses from the normal impact of tendency of many animals to choose to crowd mortality factors other than ageing. I have together in numbers in excess of 15 per nest site, omitted senescence as a mortality factor since at the peak population size of 2200 mice, 20% of rare is the animal who lives long enough in nature all nest sites were usually unoccupied. Thus there to reach a post-reproductive age without having was always opportunity for females to select an succumbed to one of the above mortality factors. unoccupied space for rearing young if they so chose. A Mortality-inhibiting Environmentfor Mice (2b) Weather amelioration: The mouse universe Some of the attributes of this environment have was located on the second floor of a prefabricated been discussed or figured elsewhere (Calhoun metal building. During the cooler months of the 1969, 1971, Wigotsky 1970). Here I shall describe year ambient temperature was kept close to 68°F how its design reduced mortality. (20°C). During the warmer months of the year large exhaust fans kept ambient temperatures (1) Emigration prevention: A closed physical mostly within the 70-90'F (21-32°C) range of the universe was formed by four 54 inch (1.37 m) outside environment. Being indoors, rain could high walls forming a square of side 101 inches not contribute to debilitation. Air movement was (2.57 m). Although the walls were structured for kept low, except when this favoured heat loss use by mice to increase the effective use-area of during periods ofhigher ambient temperature. No the universe, the mice could not climb over the evidence was ever obtained to indicate that such upper 17 inch (43 cm) unstructured portion of the weather conditions enhanced mortality.
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