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ANIMALS] fertilized zygote proceeds to form the embryo (see ).

Parthenogenesis is the production of the new organism from the female gamete without previous conjugation with the male gamete, and is to be regarded as secondary to and degenerate from true sexual reproduction. Aristotle recognized that it occurred in the bee. In 1745 C. Bonnet showed that it must occur in the case of Aphides or plant-lice, in which throughout the summer there were developed a series of generations consisting entirely of females. R. A. F. de Réaumur repeated the observations, but evaded the difficulty by suggesting that the Aphides were hermaphrodite, an explanation soon afterwards disproved by L. Dufour. In 1849 (Sir) R. Owen brought together the facts as they were then known and made a remarkable suggestion regarding them. “Not all the progeny of the primary impregnated germ cell are required for the formation of the body in all animals; certain of the derivative germ cells may remain unchanged and become included in that body which has been composed of their metamorphosed and diversely combined or confluent brethren; so included, any derivative germ cell or the nucleus of such may begin and repeat the same processes of growth by imbibition, and of propagation by spontaneous fission, as those to which itself owed its origin.” Taking hold of the recently published views of J. J. S. Steenstrup on alternation of generations, he correlated the sexual and asexual alternation in hydroids and so forth with the virgin births of insects and Crustacea, and regarded the one and the other as instances of the subsequent proliferation of included germ cells, applying the word parthenogenesis to the phenomenon. His theory was a very remarkable anticipation of the germ-plasm theory of A. Weismann, but further knowledge showed that there was an important distinction between the reproduction of the asexual generations described by Steenstrup and the cases of Aphides and Crustacea, the germinal cells in the latter instances being true ova produced from the ovaries of true females, but capable of development without fertilization. In 1856 C. T. E. von Siebold established this fact and limited Owen's term parthenogenesis to the sense in which it is now used, the development without fertilization of ova produced in ovaries. True parthenogenesis occurs frequently amongst Rotifers, and in certain cases (Philodinadae) males either do not exist or are so rare that they have not been discovered. Amongst Crustaceans it is common in Branchiopods and Ostracods; in the case of Daphnids, large thick-shelled ova are produced towards winter, which develop only after fertilization and produce females; the latter, throughout summer, produce thin-shelled ova which do not require fertilization, and from which towards autumn both males and females are produced. Amongst insects it occurs in many forms in many different groups, sometimes occasional, sometimes as a regular occurrence. Apart from Aphides the classical instance is that of the bee, where eggs that are not fertilized develop parthenogenetically and produce only drones. What is known as pathological parthenogenesis has been observed occasionally in higher animals, e.g. the frog, the fowl and certain mammals, whilst in the case of human beings, ovarian cysts in which hair and other structures are produced have been attributed to the incomplete development of parthenogenetic ova. Finally, it has been shown in a number of different instances, notably by J. Loeb, that artificial parthenogenesis may be induced by various mechanical and chemical stimulations. It has been shown that ova may be induced to segment by the presence of spermatozoa belonging even to different classes of the animal kingdom—as, for instance, the ova of echinoderms by the spermatozoa of molluscs. In such cases the resulting embryos have purely maternal characters. A possible interpretation is that spermatozoa have two functions which may be exercised independently; they may act as stimulants to the ovum to segment, and they may convey the paternal qualities. The former function may be replaced by the chemical substances employed in producing artificial parthenogenesis. Juvenile or precocious parthenogenesis, in which there takes place reproduction without fertilization in immature larvae, has been observed chiefly in insects (Dipterous midges), and to this the term paedogenesis has been applied.

The theory of parthenogenesis remains doubtful. When Weismann and others began to study the polar bodies, they made the remarkable discovery that in some parthenogenetic eggs only one polar body was extruded, but the meaning of this distinction was blurred when other cases were described in which two polar bodies were formed. Later on, Weismann drew attention to the difference between normal and reducing divisions, and it now appears to be clear that, with one set of exceptions, ova which develop without fertilization are those in which no reducing division takes place and which, accordingly, contain the number of chromosomes normal to the tissue cells of the species. Such eggs, in fact, resemble the zygote except that all their chromosomes are of maternal origin and the centrosome which becomes active in the first segmentation is that of the ovum and not, as in normal fertilized eggs, that which came in with the spermatozoon. The case of the bee and other insects in which parthenogenetic development results in the production of males, is doubtful; it appears to be the case that a reduction division has taken place in the maturation of the egg. A. Petrunkévitch has made the ingenious suggestion, that after the reducing division the normal number of chromosomes is restored by the splitting of each into two. Cases of pathological and artificial parthenogenesis would fall into line, on the supposition that the stimulus acted by preventing the occurrence of a reducing division in an ovum otherwise mature. It is to be noticed, however, that such explanations of parthenogenesis are not much more than a formal harmonizing of the behaviour of the chromosomes in the respective cases of fertilized and parthenogenetic development; they do not provide a theory as to why the process occurs.

Accessory Reproductive Organs and Processes.—It has been already stated that the primary organs of reproduction in animals are the germinal tissues producing respectively spermatozoa and ova, and that in most cases these are aggregated to form testes and ovaries. In certain animals there are no accessory organs, and when the reproductive products are ripe, they are discharged directly to the exterior if the gonads are external, as in some Coelentera, or if they are internal, break through into some cavity of the body and escape by rupture of the body-wall or through some natural aperture. In a majority of cases, however, special ducts are developed, which in the male serve primarily for the escape of the spermatozoa, but secondarily may be associated with intromittent organs. Similarly, in the female, the primary function of the gonad ducts is to provide a passage for the ova, but in many cases they serve also for the reception of spermatozoa, for the development of embryos and for the subsequent exit of the young. Associated with the ovary and the oviducts are many kinds of yolk-glands and shell-glands, the function of which is to form nutritive material for the future embryo, to discharge this into or around the ovum, and to provide protective wrappings. Although, in the last resort, fertilization depends on impulses attracting the spermatozoa to the ova, probably chemical in their nature, the necessary proximity is secured in a number of ways. In many simple cases the ripe products are discharged directly into the surrounding water, and impregnation is a matter of accident highly probable because such animals discharge enormous quantities of ova and spermatozoa, are frequently sessile and live in colonies, and are mature about the same time. In other cases, as, for instance, Tunicates and many Molluscs, the spermatozoa are discharged, and, being drawn into the body of the female with the inhalent currents, there fertilize the ova. In yet a number of other cases, there is sexual congress without intromittence. The males of many fish, such as salmon, attend the females about to discharge their ova, and afterwards pour the male fluid over the liberated eggs; whilst amongst other fish the males seek out a suitable locality and prepare some kind of nest to which the female is enticed and which receives first the ova and then the milt. In many other animals, again; as for instance the frog, the male grasps the ripe female, embracing