Life cycles adjusted to environmental change
Both sexual and nonsexual reproduction may be exploited or adjusted to meet widely fluctuating environmental conditions, especially those of a regular seasonal character. This phenomenon is particularly striking in the case of the smaller or simpler forms of animal and plant life that have a life-span of a year or less. The seeds of annual plants germinate in the spring, grow and set seed in turn during the summer, and die in the fall. Only the sexually produced seeds persist and represent the species during the long winter season. Certain small, though common, freshwater creatures have a similar cycle. The microscopic eggs of Hydra and of Daphnia, for example, lie at the bottom of ponds throughout the winter, each within a tough protective case. In late winter or early spring, a new generation of hydras develops, each individual becoming attached to a stone or vegetation and feeding on small crustaceans by means of its long slender tentacles. The daphnias, or so-called water fleas, emerge at about the same time and grow rapidly to maturity. In both cases the growing season, usually from spring until fall, is a time for intensive reproduction by whatever means is most effective. Hydras bud off new hydras continually, each new hydra repeating the process, with the size of populations limited only by available food. Only late in the season, when the food supply drops off and the temperature drops, does the riotous splurge of nonsexual reproduction come to an end. Then each individual ceases to bud and produces either minute ovaries or testes, and in some species, both. Eggs become fertilized, encased, drop into the mud, and await the coming of the following spring, while the parental creatures die as living conditions worsen with approaching winter. Such is a general pattern of life, widely seen among creatures whose individual existence is measured in weeks or months but whose race must persist in some form at all times if extinction is to be avoided.
So it is with Daphnia and many other organisms. The Daphnia also changes according to the times, but it alternates between one form of sexual reproduction and another. Sexually, the Daphnia is exquisitely adapted to the little world in which it lives. Under ideal conditions every member of a Daphnia community is female. All those first hatching out from winter eggs in the spring are females. Each produces a succession of broods during the month or two of its individual existence, all offspring being females. Each such female, generation after generation, during the spring and summer seasons, produces eggs that develop at once without need or opportunity of being fertilized. No males in fact are present. Every individual is a self-sufficient breeding female. Population explosions occur wherever environmental circumstances are favourable. Eventually, however, conditions inevitably change for the worse, either because of effects inherent in any population explosion or because every season comes to an end. Food becomes scarce because of too many consumers; space becomes crowded and in some degree polluted; chilly days succeed the warmth of summer. Whatever the cause, and well before disaster can strike, the creatures respond in remarkable ways. On the first signal that conditions may be getting less than good, a certain number of the eggs produced by a population of Daphnia develop into males, each with testes in place of ovary, together with certain secondary sexual characteristics. A scattering of males through the virgin paradise, however, is only the first step, a preparedness in case conditions go from bad to worse. If there has been a false alarm, the females continue to produce female-producing eggs that develop parthenogenetically—that is, without benefit of fertilization—and the males die off without performing any sexual function. But if the environmental signal means the beginning of the end of congenial conditions, a cell in the ovary of each female grows to form a larger egg than usual, and it is of a type that must be fertilized. Then mating between the sexes takes place, and the resulting special, fertilized eggs become thickly encased and alone survive the winter season after becoming separated from the parent.
Wherever small aquatic creatures live in bodies of water that may freeze in winter or dry up in summer, similar adaptations may be seen in many forms of life besides hydras and water fleas. Certain small fish, known as the annual fishes, have individual life-spans of about six months. The life-span itself is in fact adapted to the period during which active existence is possible in their particular habitat. When the water holes, swamps, and puddles in which they live begin to dry up, mating takes place, and the fertilized eggs drop into the mud. The parents die, and the eggs remain in a state of suspended development until the next rainy season occurs. The race must continue whatever the circumstances, and all sex is directed toward this end.
The origin of sex and sexuality
All sexual reproduction, no matter how large or small the organisms may be, is a performance of single cells. Only at the level of single cells can the essential genetic recombinations be accomplished. So in every generation new life begins with the egg, which is a single cell, however large it may be. Egg and sperm unite at fertilization, but the fertilized egg is as much a single cell as before. When did it all begin? The generally accepted answer is that the fundamental, or molecular, basis of sexuality is an ancient evolutionary development that goes back almost to the beginning of life on earth, several billion years ago, for it is evident among the vast world of single-celled organisms, including bacteria.
In these lowest forms of life, sex and reproduction are distinct happenings. Reproduction is accomplished in most cases entirely by fission, which is simply cell division repeated regularly, as long as the environmental conditions permit. As long as crowding and other adverse changes are avoided, cells divide, and the daughter cells grow and divide again, for weeks or months on end. This process occurs in both plantlike and animal-like single-celled organisms and in bacteria as well. Under certain other conditions, such cell organisms come together and fuse in pairs, a form of sexual behaviour at its primary level and comparable to the fusion of an egg and sperm. In all such case, a combined cell is produced in which nuclear exchange or recombination has occurred. Pairing off of this sort takes place sooner or later in all forms of unicellular life, even where no outwardly distinguishable differences can be detected between the pairing individuals. The lack of discernible differences between the members of mating pairs, however, does not mean that pairing occurs between identical individuals. In the much investigated Paramecium and other protozoan organisms, two separate populations of cells may continue to increase almost indefinitely by ordinary cell division of single individuals, but when two such populations are mixed together, mating generally occurs immediately between individuals from the two different sources. The fusion, or pairing, has essentially the same function as the fusion of the male and female nucleus during the process of fertilization of eggs of higher forms. It is the basis of sex, the essential event in all cases being the genetic or chromosomal recombination.
Individual mating cells (i.e., eggs, sperm, or even whole single-celled organisms) may be called gametes whether or not they are distinguishable from one another. Yet even among the varius single-celled organisms, mating commonly occurs between individuals of two different kinds. This kind of mating is seen most often among the single-celled organisms known as flagellates. In some species the gametes may be alike and all are motile, progressing through the water by means of one or more whiplike flagella similar to the tail of a sperm. In other species, all individuals may still be motile, but pairing occurs between individuals of different sizes. In still others, one of the two mating types may be very small and motile, and the other, large, with stored nutritional material, and nonmotile. All degrees of differentiation between male and female gametes can be found, and it is probable that the basic and characteristic distinction between the sex cells of both animal and plant life in general was established very early in the course of evolution, during the immense period of time when virtually all living organisms consisted of single cells.
This division of labour between mating types, male and female, respectively, is nature’s way of attaining two ends. These are the bringing together of the gametes so that fusion may take place and the accumulation of reserves so that development of a new organism can be accomplished. The first calls for as many motile cells as possible; the second calls for cells as large as possible. These different requirements are practically impossible to satisfy by a single type of cell. Accordingly, and especially in multicelled animals of all sorts, male gametes, or spermatozoa, are extremely small, extremely motile, and are produced in enormous numbers. The larger the number, the greater the likelihood that some will encounter and fertilize eggs. On the other hand, the female gametes, or ova, individually need to be as large as possible since the larger the size and the more condensed the internal nutritional reserves, the farther along the path of embryonic development the egg can travel before hatching must occur and the new organism must fend for itself. Nevertheless, eggs in general are caught between the desirability of being individually as large as they can be and the persisting need to be produced in reasonably large numbers, so that an assortment of differing individuals is produced from a single pair of parents. A large number of offspring ensures that a proportion, at least, will survive the environmental hazards faced by all developing organisms in some degree.