that each spermatocyte gives origin to four spermatids. Each
spermatid becomes a functional spermatozoon or male gamete.
The gametogeny of the male therefore closely resembles that of the
female, differing from it only in the fact that all the four products
of the progamete become functional gametes, whereas in the female
only one, the ovum, becomes functional, the other three (polar bodies)
being abortive. In the spermatogenesis of the bee, however, the
spermatocyte only divides once, giving rise to a small polar-body-like
structure and one spermatid (Meves, Anat. Anzeiger, 24, 1904, pp.
29-32). The nucleus of the male gamete is not called the male pronucleus,
as would be expected, that term being reserved for the
second nucleus which appears in the ovum after fertilization. As
this is in all probability derived entirely from the nucleus of the
spermatozoon, we should be almost justified in calling the nucleus
of the spermatozoon the male pronucleus. In most forms in which
the formation of the gametes from the progamete has been accurately
followed, and in which the progamete of both sexes divides twice in
forming the gametes, the division of the nucleus presents certain
peculiarities. In the first place, between the first division and the
second it does not enter into the resting state, but immediately
proceeds to the second division. In the second place, the number of
chromosomes which appear in the final divisions of the progametes
and assist in constituting the nuclei of the gametes is half the number
which go to constitute the new nuclei in the ordinary nuclear divisions
of the animal. The number of chromosomes of the nucleus of the
gamete is therefore reduced, and the divisions by which the gametes
arise from the progametes are called reducing (maiotic) divisions.
It is not certain, however, that this phenomenon is of universal
occurrence, or has the significance which is ordinarily attributed to it.
In the parthenogenetic ova of certain insects, e.g. Rhodites rosae
(Henking), Nematus lacteus (Doncaster, Quart. Journal Mic. Science,
49, 1906, pp. 561-589), reduction does not occur, though two polar
bodies are formed.
As soon as the spermatozoon has conjugated with the ovum, a second nucleus appears in the ovum. This is undoubtedly derived from the spermatozoon, possibly from its nucleus only, and is called the male pronucleus. It possesses in the adjacent protoplasm a well-marked centrosome. The Fertilization. general rule appears to be that the female pronucleus is without a centrosome, and that no centrosome appears in the female in the divisions by which the gamete arises from the progamete. If this is true, the centrosome of the zygote nucleus must be entirely derived from that of the male pronucleus. This accounts for the fact, which has been often observed, that the female pronucleus is not surrounded by protoplasmic radiations, whereas such radiations are present round the male pronucleus in its approach to the female. In the mouse the subsequent events are as follow:—Both pronuclei assume the resting form, the chromatin being distributed over the nuclear network, and the nuclei come to lie side by side in the centre of the egg. A long loop of chromatin then appears in each nucleus and divides up into twelve pieces, the chromosomes. The centrosome now divides, the membranes of both nuclei disappear, and a spindle is formed. The twenty-four chromosomes arrange themselves at the centre of this spindle and split longitudinally, so that forty-eight chromosomes are formed. Twenty-four of these, twelve male and twelve female, as it is supposed, travel to each pole of the spindle and assist in giving rise to the two nuclei. At the next nuclear division twenty-four chromosomes appear in each nucleus, each of which divides longitudinally; and so in all subsequent divisions. The fusion of the two pronuclei is sometimes effected in a manner slightly different from that described for the mouse. In Echinus, for instance, the two pronuclei fuse, and the spindle and chromosomes are formed from the zygote nucleus, whereas in the mouse the two pronuclei retain their distinctness during the formation of the chromosomes. There appears, however, to be some variation in this respect: cases have been observed in the mouse in which fusion of the pronuclei occurs before the separation of the chromosomes.
Parthenogenesis, or development of the female gamete without fertilization, is known to occur in many groups of the animal kingdom. Attempts have been made to connect this phenomenon with peculiarities in the gametogeny. For instance, it has been said that parthenogenetic ova Parthenogenesis. form only one polar body. But, as we have seen, this is sometimes the case in eggs which are fertilized, and parthenogenetic ova are known which form two polar bodies, e.g. ova of the honey-bee which produce drones (Morph. Jahrb. xv., 1889, p. 85). ova of Rotifera which produce males (Zool. Anzeiger, xx., 1897, p. 455), ova of some saw-flies and gall flies which produce females (L. Doncaster, Quart. Journ. Mic. Sc., 49, 1906, pp. 561-589). Again it has been asserted that in parthenogenetic eggs the polar bodies are not extruded from the ovum; in such cases, though the nucleus divides, those of its products which would in other cases be extruded in polar bodies remain in the protoplasm of the ovum. But this is not a universal rule, for in some cases of parthenogenesis polar bodies are extruded in the usual way (Aphis, some Lepidoptera), and in some fertilized eggs the polar bodies are retained in the ovum.
It is quite probable that parthenogenesis is more common than has been supposed, and it appears that there is some evidence to show that ova, which in normal conditions are incapable of developing without fertilization, may yet develop if subjected to an altered environment. For instance, it has been asserted that the addition of a certain quantity of chloride of magnesium and other substances to sea-water will cause the unfertilized ova of certain marine animals (Arbacia, Chaetopterus) to develop (J. Loeb, American Journal of Physiology, ix., 1901, p. 423); and according to M. Y. Delage (Comptes rendus, 135, 1902. Nos. 15 and 16) such development may occur after the formation of polar bodies, the chromosomes undergoing reduction and the full number being regained in the segmenting stage. These experiments, if authenticated, suggest that ova have the power of development, but are not able to exercise it in their normal surroundings. There is reason to believe that the same assertion may be made of spermatozoa. Phenomena of the nature of parthenogenesis have never been observed in the male gamete, but it has been suggested by A. Giard (Cinquantenaire de la Soc. de Biol., 1900) that the phenomenon of the so-called fertilization of an enucleated ovum which has been described by T. Boveri and Delage in various eggs, and which results in development up to the larval form (merogony), is in reality a case in which the male gamete, unable to undergo development in ordinary circumstances on account of its small size and specialization of structure has obtained a nutritive environment which enables it to display its latent power of development. Moreover, A. M. Giard suggests that in some cases of apparently normal fertilization one of the pronuclei may degenerate, the resultant embryo being the product of one pronucleus only. In this way he explains certain cases of hybridization in which the paternal (rarely the maternal) type is exclusively reproduced. For instance, in the batrachiate Amphibia, Héron Royer succeeded in 1883 in rearing, out of a vast number of attempts, a few hybrids between a female Pelobates fuscus and a male Rana fusca; the product was a Rana fusca. He also crossed a female Bufo vulgaris with a male Bufo calamita; in the few cases which reached maturity the product was obviously a Bufo calamita. Finally, H. E. Ziegler (Arch. f. Ent.-Mech., 1898, p. 249) divided the just-fertilized ovum of a sea-urchin in such a way that each half had one pronucleus; the half with the male pronucleus segmented and formed a blastula, the other degenerated. It is said that in a few species of animals males do not occur, and that parthenogenesis is the sole means of reproduction (a species of Ostracoda among Crustacea; species of Tenthredinidae, Cynipidae and Coccidae among Insecta); this is the thelytoky of K. T. E. von Siebold. The number of species in which males are unknown is constantly decreasing, and it is quite possible that the phenomenon does not exist. Parthenogenesis, however, is undoubtedly of frequent occurrence, and is of four kinds, namely, (1) that in which males alone are produced, e.g. honey-bees (arrhenotoky); (2) that in which females only are produced (thelytoky), as in some saw-flies; (3) that in which both sexes are produced (deuterotoky), as in some saw-flies; (4) that in which there is an alternation of sexual and parthenogenetic generations, as in Aphidae, many Cynipidae, &c. It would appear that “parthenogenesis does not favour the production of one sex more than another, but it is clear that it decidedly favours the production of a brood that is entirely of one sex, but which sex that is differs according to circumstances” (D. Sharp, Cambridge Natural History, “Insects,” pt. i. p. 498). In some Insecta and Crustacea exceptional parthenogenesis occurs: a certain proportion of the eggs laid are capable of undergoing either the whole or a part of development parthenogenetically, e.g. Bombyx mori, &c. (A. Brauer, Arch. f. mikr. Anat., 1893; consult also E. Maupas on parthenogenesis of Rotifera, Comp. rend., 1889–1891, and R. Lauterborn, Biol. Centralblatt, xviii., 1898, p. 173).
The question of the determination of sex may be alluded to
here. Is sex determined at the act of conjugation of the two
gametes? Is it, in other words, an unalterable property
of the zygote, a genetic character? Or does it depend
upon the conditions to which the zygote is subjected in
Determination
of sex.
its development? In other words, is it an acquired
character? It is impossible in the present state of knowledge to
answer these questions satisfactorily, but the balance of evidence
appears to favour the view that sex is an unalterable, inborn character.
Thus those twins which are believed to come from a split zygote
are always of the same sex, members of the same litter which have
been submitted to exactly similar conditions are of different sexes,
and all attempts to determine the sex of offspring in the higher
animals by treatment have failed. On the other hand, the male
bee is a portion of a female zygote—the queen-bee. The same
remark applies to the male Rotifer, in which the zygote always
gives rise to a female, from which the male arises parthenogenetically,
but in these cases it does not appear that the production of males
is in any way affected by external conditions (see R. C. Punnett,
Proc. Royal Soc., 78 B, 1906, p. 223). It is said that in human
societies the number of males born increases after wars and famines,
but this, if true, is probably due to an affection of the gametes
and not of the young zygote. For a review of the whole subject
see L. Cuénot, Bull. sci. France et Belgique, xxxii., 1899, pp. 462-535.
The first change the zygote undergoes in all animals is what is generally called the segmentation or cleavage of the ovum. This consists essentially of the division of the nucleus into a number of nuclei, around which the protoplasm sooner or later
