SEX (Lat. sexus; possibly connected with secare, to cut), the character of being either male or female, which can be attributed to the vast majority of animals, but less correctly to the higher plants, where the so-called male and female organs, or flowers, are part of the sexless generation (see Reproduction: Plants). The primary distinction of sex resides in the essential organs of reproduction (q.v.). An organism that contains the germinal tissue or mass of tissue known as the testis, and producing the sexual cells known as spermatozoa, is a male; an organism containing the tissue which produces ova is known as a female; one producing both ova and spermatozoa is a true hermaphrodite; and one producing neither, if it belong to the sexual generation, is known as a neuter, although neuters are for the most part incomplete females. The primary sexual tissues and the gametes are described in the article Reproduction (Animals).
Associated with the presence of the primary reproductive organs there may be a large number of other characters, and attempts have been made to classify these as secondary and tertiary sexual characters. It is impossible to define a series of logical categories in which any accessory character will find its inevitable place, but a convenient practical distinction first made by John Hunter may be drawn between characters directly auxiliary to the processes of reproduction and those which, although limited to one sex, are not immediately connected with reproductive processes. We may then make the division into (1) Primary Sexual Characters (A. Essential: power of producing respectively ova and spermatozoa. B. Auxiliary: possession of sexual ducts and reservoirs, intromittent and copulatory organs, organs associated with oviposition, gestation, parturition, and nutrition of the immature young in any stage); and (2) Secondary Sexual Characters (differences between the sexes in size, shape, appearance, ornamentation, armament, colour and coloration, voice, and instincts and habits not directly associated with the reproductive processes).
Those characters which are here grouped as primary are described in the article Reproduction. It is sufficient to repeat that in many animals only the essential primary characters are present. There is much diversity in the possession of secondary sexual characters, and in many cases these apparently are absent. Among mammals it is impossible to distinguish the sex without examination of the reproductive organs or observation of the sexual habits, in such cases as the domestic cat, the tiger and many other feline animals, hyaenas, bears, rabbits, hares, mice and a vast number of others. So also among birds there are many cases where the sexes are alike, as for instance, some hummingbirds, parrots, owls, cranes, kingfishers, and many small birds such as robins and hedge-warblers. In reptiles and batrachians, in fish and a very large number of invertebrates there are no visible secondary sexual characters.
C. Darwin, in the portion of the Descent of Man devoted to “Selection in relation to Sex,” brought together what remains the most complete and valuable account of the existence and distribution amongst animals of secondary sexual characters, and it would be impracticable here to give more than the most summary description of the groups of facts involved. Among Crustacea the sexes frequently differ, but in most cases the differences concern auxiliary primary characters, such as the possession of intromittent and clasping organs. Differences in size are frequent; in the higher Decapods the males and in the lower Crustacea the females frequently being larger, the disparity being extreme in some of the parasitic Copepods and Isopods where the males are minute and attached to the females, whilst in the Cirripedes, as Darwin himself discovered, very minute complementary males may live as parasites in the mantle cavity of large hermaphrodite or female forms. Amongst Arachnids conspicuous differences in colour and size occur, the males generally being smaller, more active and possessed of relatively longer appendages, and more highly decorated. Amongst Insects, the differences between the sexes may be very great, quite apart from those relating to intromittence, pretension of the female, oviposition, or the higher development of sense organs by which the males can more readily seek out the females. In many cases the males are winged, the females wingless and grub-like. In a few instances, the males are highly pugnacious and are furnished with special weapons for fighting with their rivals. Amongst the Homoptera and Orthoptera there are many instances where the males possess organs capable of producing loud sounds, and these are rudimentary or absent in the females, whilst in other cases, both sexes produce call-notes. Particularly amongst the Coleoptera, the males may differ very greatly from the females in the shape of the body and may be decorated with extraordinary growths of the head and thorax. The most notable sexual differences are in coloration, and whilst there are many instances where both sexes are inconspicuous, and a few where both are brilliant, there are still more where the males differ from the females by the display of more conspicuous patterns and of brighter colours. It may be said of Insects in general that it is the more common case for secondary sexual characters to exist in such a degree that the sexes may be distinguished at a glance.
Among Fishes, secondary sexual characters are common. Spines are developed on the head and pectoral fins of the males of some Rays, but it is probable that these may be auxiliary primary characters, useful in the pretension of the female. In the male salmon, a cartilaginous projection, developed during the breeding season, appears on the upper surface of the point of the lower jaw, whilst in old males the jaws become hook-like and the teeth are greatly increased in size. In the thornback, the adult male has the teeth sharp-pointed and backwardly-directed, while those of the female are flat and pavement-like. In almost all fishes the males when adult are smaller than the females, and may be much smaller. Beards of stiff, hair-like structures, elongated processes of the fins, tubercles and many other structures that may be classed as ornaments, because their function is unknown, occur in males and are absent in females. Differences in pattern and colour are extremely frequent, become much more marked in the breeding season, and are of such a nature that the males are more conspicuous. Among Batrachia, differences between the sexes in size and general shape are not striking, but there are many instances of the males exhibiting crests, or special processes which may be classed as ornaments, and peculiar patterns and bright colours, during the breeding season.
Secondary sexual differences appear in the vast majority of birds. The shape seldom differs markedly, but differences in size are common, sometimes, as in birds of prey, the females, and sometimes, as in the allies of the domestic fowl, the males being larger. In a large number of instances the males are very pugnacious and are better armed, the bones and musculature being heavier, the beaks and claws stronger, while spurs or knobs on the wings and spurs on the legs may be present only in the males or be relatively small in the females. Special ornaments such as crests and wattles, combs, carbuncles, excrescences of the skin, and elongated or peculiarly shaped feathers are extremely frequent, and are developed or intensified in the breeding season, and in the vast majority of cases confined to the males. The voice almost invariably varies with the sex, is associated with the breeding period and is much more highly developed in the male, whilst structural developments such as modifications of the trachea, vocal sacs and resonators and differences in the larynx are frequently present and on the whole distinctive of the males. Differences in colour and pattern are extremely well marked, and these are well known to be associated with the breeding period, which in many cases is preceded by a moult, after which the sexual plumage is assumed, or the colour of the naked parts intensified. In a few exceptional cases such as some button-quails (Turnix), painted snipes (Rhynchaea), phalaropes (Phalaropus), and cassowaries, the females exceed the males in size and brilliancy, and it is interesting to notice that in such cases the usual distinction of habit may be reversed, the females being pugnacious, aggressive, and courtiers of the males, whilst the latter are shy and may attend to the brood. Such exceptions are so rare that they may be called abnormal, for the rule among birds is that where secondary sexual characters are displayed, ornamentation, voice, brilliant pattern and colour, pugnacity and amorousness are distinctive of the male. Secondary sexual differences of the same nature are abundant among mammals. The males are usually larger and have greater strength with corresponding bones and muscles, and courage and pugnacity. Special weapons of offence or defence are common and are usually limited to the males or more highly developed in them; familiar instances are the horns of cattle, sheep and antelopes, the canine teeth, the mane of the lion. The antlers of the stags are certainly used in combats between the males, but in their more extreme development they may be classed as sexual ornaments. The males of many mammals emit powerful odours during the breeding season, whilst their voices, whether as a battle cry or a call to the female, are frequently more powerful. Crests, tufts and mantles, rudimentary in the female, conspicuous in the male, are extremely common. Differences in pattern and colour are rare except in monkeys, but when these exist they are usually found in the male.
The sexes, then, are distinguished by primary and secondary characters, these two categories being convenient rather than logical. The real dividing line is between the essential primary sexual character, the presence of a male or female gonad, and the various auxiliary and secondary differences which appear in every grade of elaboration. It is to be noted, moreover, that all the other sexual characters depend on the activity of the essential primary character. Immature males and females are closely alike; the auxiliary and secondary sexual characters almost invariably begin to appear only when the gonads become mature, and fade away when these are injured or destroyed by accident, disease, senescence or artificial interference, and finally, when the activity of the gonads waxes and wanes periodically, there is a corresponding periodicity in the display of the secondary characters. A number of observations and experiments support the conclusion that the gonads, in addition to their obvious function of producing the sexual cells, discharge secretions into the blood and tissues, and that these internal secretions or hormones, are the physiological stimulus which awakens the development of the auxiliary and secondary sexual characters.
Auxiliary primary and secondary sexual characters are so many and various that general statements regarding them are difficult and uncertain. In the broadest fashion, however, the following generalizations appear to be true. Secondary sexual characters begin to appear at puberty. Young or immature forms resemble the sex in which such characters are least marked, while the young and the undistinguished sex resemble ancestral forms. The sex that is distinguished is usually the male, and the characters are usually hypertrophies or specializations of characters that appear in the females and the young. (It is to be remembered that specialization may be the result of the suppression of characters as well as their acquisition, and there are a remarkable number of cases in which we may, at least tentatively, picture the bright sexual colour of males as due to the suppression of a pigment which masks them in the female.)
Hermaphroditism is the condition in which gonads producing ova and gonads producing spermatozoa are contained in the same individual. Its distribution in the animal kingdom is irregular, and apparently independent of natural affinity, and the balance of opinion is in favour of regarding it not as a primitive condition, but as a secondary acquisition. C. Claus has pointed out that it is frequent among sessile animals, as for instance Sponges, Anemones, Corals, Polyzoa, bivalve Molluscs, and Tunicates, and sluggish animals such as many of the worms and snails, whilst it is extremely common amongst almost every kind of parasitic animal. The obvious suggestion is that if the condition be primitive, it has been preserved, and if not primitive, acquired, because in animals of such habit, the chances of sexual congress would be greater than if the sexes were separate. Against such an interpretation, however, it must be noticed that in most hermaphrodites the sexual maturity of the male and female gonads is not coincident, so that cross-fertilization commonly occurs. Self-fertilization is said to occur in the fish Serranus, and it certainly occurs in many parasitic Trematodes, in Tapeworms and a few Nematodes. The real meaning of the occurrence of the condition remains obscure. Both gonads are present in many Sponges, in the Ctenophora, in many Anemones and Corals, in degenerate Hydroids such as Hydra, in most Turbellarians and Trematodes, in all the Tapeworms, in a few Nematodes, in many Chaetopods, in the Leeches, in a few Brachiopods and in many Polyzoa. It is absent in most Echinoderma and Arthropoda, but occurs in Cirripedes and some Isopods. It occurs in some bivalves, such as the common oyster, cockle and clam, and is present in the Euthyneurous Gastropods and in Pteropods. Amongst vertebrates it is rare. A number of observers have urged that the vertebrate embryo passes through a hermaphrodite condition. J. T. Cunningham and F. Nansen have stated that a testis is embedded in the ovary of the young hagfish (Myxine) and that this ripens before the ovary, but later observers have disputed their interpretation of the facts. In a few fish and some Batrachia, hermaphroditism has been demonstrated, but it is not certain, whether as a normal or aberrant occurrence, whilst in many of the Batrachian cases, the animals are known to be normally unisexual. The term hermaphroditism, however, has been applied frequently to cases of a different kind, in which there is no evidence of the essential sexual organs being affected, the appearances relating wholly to the auxiliary primary or the secondary sexual characters. It is most probable that such conditions differ entirely from true hermaphroditism. With regard to the auxiliary primary organs, and especially the genital ducts and external organs of sex, in a majority of cases as in vertebrates, the embryonic or youthful condition is undifferentiated, and so to say, contains the initial material which may be elaborated by specialization in one direction or the other, by the proliferation of certain portions and the suppression of others, into the structures characteristic of the male or of the female. Sometimes, growth takes place without normal differentiation, sometimes the specialization in one direction lags, with the result that a dubious appearance arises. Subsequent dissection or the approach of maturity, however, make it plain that the dubiety was superficial and that the gonad of only one sex was present. Among mammals, including man, every normal male retains relics of the female side of the undifferentiated condition of the accessory sexual organs, whilst every normal female contains similar if less well-marked relics of the male condition. Apparent hermaphroditism depending on a dubious condition of the secondary sexual characters is equally widespread in possible occurrence. Amongst insects which have been much studied, such as the butterflies and moths, many curious conditions have been described; sometimes the pattern and colour of the upper and under sides, sometimes of different parts of the same wing, sometimes of different wings, present the characters of different sexes. Among birds and mammals, the secondary sexual characters of one sex, such as size, pattern or colour, weapons or habits, may appear in animals with the gonads of the other sex, in every degree of development, reaching to an apparently complete reversal. In many cases these abnormal occurrences are associated with arrest of the functional activity of the primary organs of sex, by disease, accident, or decay, and the failure of the necessary stimulus would certainly serve to explain cases where the apparent reversal is no more than the suppression of a specialization in one direction. The facts, however, go further; it appears as if the suppression of femaleness allows the development of a latent maleness.
Determination of Sex.—Answers to the question why a particular individual becomes a male or a female fall into two groups, in one of which it is supposed that external conditions determine the result, in the other that the sexual cells differ from the first. G. Canestrini suggested that the sex was determined by the number of spermatozoa which entered the ovum, but fuller knowledge of the details of fertilization (see Reproduction) has made it plain that only a single spermatozoon, normally conjugates with the ovum, whilst polyspermy, if it occur, results only in abnormalities which do not proceed to full development. Professor Thury in 1863 and C. Düsing in 1883 urged that ova fertilized soon after ovulation gave rise to females, whilst those impregnated later produced males. Some evidence exists as to the effect of delay in fertilization; V. Hensen (1881) suggested that females were produced when both ova and spermatozoa were in the most active condition, and H. M. Vernon (1898) has shown that in hybridizing Echinoderms the fresher gamete appears to exert a greater influence, but it cannot be said that there is definite evidence as to the determination of sex on such lines. J. D. Hofacker in 1823 and M. T. Sadler in 1830 collected a large series of statistics from which they drew the conclusion that when the male parent is older, more males are produced, whilst many observers have attempted to draw conclusions from the comparative vigour of the parents. Popular belief and some observations with regard to the breeding of domestic animals have led to the inference that the sex of the offspring tends to be that of the least vigorous parent, and such a theory, as it would appear to imply the existence of a natural law for rectifying the proportions of the sexes, has gained more attention than the facts supporting it would justify, and several unbiased observers have interpreted the events in the sense that the vigorous parent produces his or her own sex. It is to be noted that such theories of relative vigour do not necessarily imply that external conditions determine the sex, for they would apply equally were it the case that there was a power of selection amongst gametes of predetermined sex. A large number of investigators have been led to believe that conditions of nutrition are of importance, and this view is specially plausible in the case of vertebrates, if it be accepted that the embryos pass through a hermaphrodite condition. E. Yung found that when tadpoles were reared under normal conditions, the proportion of male to female was about as 43 to 57, but that when a flesh diet was provided the percentage of females was very greatly increased. It has been noted that when Aphides are under the favourable conditions of summer temperature and nutrition, they produce only females, but that the advent of autumn brings with it an equality in sex production. Mrs Treat showed that starved caterpillars turned into males; E. Maupas, in the case of Rotifers, and other observers in the cases of some Crustacea, have similarly pointed to a relation between abundant nutrition and the excessive production of females. In nearly every case, however, other observers have either obtained conflicting results, or placed another interpretation on similar results, whilst in none of the cases has the factor of selective mortality been sufficiently excluded. Even were it proved that a correlation existed between excessive diet and over-production of females, it might be that the incidence of mortality was differential. Many attempts have been made to derive information by examining the statistics of human births in times of plenty and of hardship, but the results are inconclusive. C. Darwin, reviewing the evidence was disposed to believe that the proportions of the sexes varied, that the tendency to produce male and female offspring was inherited, and that by a process of natural selection it was adjusted to the needs of the species, but he was too cautious to lean to any particular view as to the nature of the determining factors. C. Düsing (1883 and 1885) also believed in the existence of such a power of adaptation or adjustment, and attributed it to the action of a large number of external conditions. P. Geddes and J. A. Thomson (1889) similarly came to the conclusion that factors external to the sexual cells had a predominating importance, and these authors linked the determination of sex with their general theory of the nature of sex. They regarded sex as an expression of an alternating rhythm of anabolism and katabolism to be observed throughout the living world, and supposed that femaleness was specially associated, was in fact an outcrop of the anabolic or constructive processes of living matter, whilst maleness represented the katabolic, destructive or liberating processes. Their view ranges many diverse facts in apparent harmony, but has to encounter many facts that apparently contradict it. In a later work J. A. Thomson himself (1907) assigns less weight to his own theory, and quotes with approval T. H. Morgan's suggestion that the determination of sex may be brought about in different fashions in different cases.
Theories as to sex being predetermined in the sexual cells have been numerous, but it is only recently that any exact evidence appearing to point to such a conclusion has been adduced. When parthenogenesis (see Reproduction) was first being investigated, it was found that eggs which gave rise to females were different from those which produced males, but when it was demonstrated that at least in many cases there was the further difference as to whether the eggs were fertilized or not, it was assumed that the presence or absence of fertilization determined the sex. Physicians have repeatedly propounded the theory that one ovary produces eggs capable of developing only into females, the other only those capable of becoming males, and the suggestion has been made that in the case of human beings ovulation takes place alternately from the ovaries. From this it would follow that were the sex resulting from one fertilization known, the sex of a subsequent fertilization could be predicted, or by choosing the date of fertilization, selected. These views, however, rest on no satisfactory evidence and remain uncorrelated with any observations as to the structure of the eggs themselves. On the other hand, more exact workers, using modern cytological methods, have accumulated striking facts as to the existence of different kinds of sexual cells, the differences relating chiefly to the nuclear changes which occur in ovogenesis and spermatogenesis, and have been established with more certainty in the case of the spermatozoa. E. B. Wilson (1909) has given a full summary and discussion of various interpretations of these observations. In over a hundred species of insects, Myriapods and Arachnids, two kinds of spermatozoa are produced. The spermatozoa are formed in pairs, and the mother cell which gives rise to each pair exhibits, in the ordinary fashion of nuclear division, paired chromosomes, one member of each pair passing into each spermatozoon. The mother cell contains also an unpaired element, consisting in its simplest form of a single large chromosome, but sometimes represented by a group of peculiar chromosomes, which, for convenience, Wilson terms the “X” element, or “heterochromosome." The “X” element passes into one or other of the spermatozoa, from which it results that spermatozoa of two kinds are formed in equal numbers, the difference being the presence or absence of the “X” element. Eggs fertilized by spermatozoa containing the “X” element become females, those fertilized by spermatozoa without it become males. There is evidence that in some cases (e.g. bees) the spermatozoa devoid of the “X” element degenerate, with the result that any fertilized eggs must produce females.
E. B. Wilson's suggestion, advanced in the most cautious way, is that the “X” element referred to in the last paragraph is the determinant, or at least the index, of sex, and further that the difference between the male and female organism is that the male comes from an egg which, developing either parthenogenetically or after fertilization, contains only a single unit of the “X” element, while the female starts from an ovum which, whether developing after fertilization or parthenogenetically, contains the two “X” units. The ovum of a sexual egg in the process of maturation discards half its normal complement of the “X” element; if it be fertilized by a spermatozoon containing an “X” unit it gives rise to a female; if it be fertilized by one without this it becomes a male. A large number of different forms of nuclear change have been described in the maturation of normal and parthenogenetic eggs, and by the exercise of a little ingenuity it is easy to select from these various processes modes of nuclear division which if they actually occurred in the appropriate instances would adapt Wilson's hypothesis to cases in which parthenogenetic eggs give rise to males or to females. In some individual instances the process which the hypothesis would demand appears actually to occur.
Various workers on Mendelian lines (see Mendelism) have endeavoured to correlate the facts discussed by Wilson and their experimental inquiries into the inheritance of primary and secondary sexual characters, with the additional difficulty, absent from Wilson's hypothesis, that their theory requires them to suppose the unfertilized cells to be unisexual. W. E. Castle suggested that both males and females were Mendelian male-female hybrids with respectively male and female dominance, and that in the usual way disruption took place in the formation of the germ cells, with the result that male and female spermatozoa and male and female ova were produced. He assumed further that there was a selection or repulsion in fertilization, so that ova and spermatozoa bearing the same sex never conjugated. C. Correns assumed the male to be sex-hybrid, the female to be homozygous or pure female, the male character being dominant. Ova were, therefore, unisexual, always female, while spermatozoa were either male or female, and when a female egg was fertilized by a female spermatozoon the result naturally was a female, but when it was fertilized by a male spermatozoon the result was a sex-hybrid appearing as a male because of the dominance of male characters. Correns's theory avoids the unlikely supposition of selective fertilization, but breaks down in those cases of parthenogenesis where the unfertilized egg produced by a female gives rise to a male. W. Bateson reverses the theory of Correns and supposes that the female is a hybrid with femaleness dominant, while the male is pure male. The female in fact contains a factor which makes her female whilst the male is a male because it is without this factor. This view, however, leaves unexplained the existence of two kinds of spermatozoa and involves a series of elaborate hypotheses to reconcile it with cases of parthenogenesis. L. Doncaster has elaborated the extremely ingenious suggestion that the Mendelian pairs are not male and female, but male and absence of sex and female and absence of sex. The male is a pure male but produces two kinds of spermatozoa, those with the determinant for sex and those without it. The normal female is a sex-hybrid and produces male and female eggs in equal numbers, and it is assumed that there is a selective fertilization, female eggs being fertilized by male spermatozoa and giving rise to females, whilst male eggs are fertilized by spermatozoa without the sex factor and give rise to males. In cases of parthenogenesis, it is supposed that there are two kinds of females, the result of fertilization by different kinds of spermatozoa, and that those going through different kinds of maturation processes give rise without fertilization to males or to females. Doncaster has discovered many interesting details of the maturation processes in insects which agree with his suggestion. The Mendelian interpretations, however, are more ingenious than conclusive, but at least they combine with other work in supporting the probability that the determination of sex depends on the sexual cells and not on conditions influencing the developing embryo. Similarly they combine with other work in pointing to the conclusion that the male organism differs from the female by the absence of something present in the female. The Mendelian interpretations suggest that male and female sex determinants are different in kind; Wilson's interpretation suggests that they differ only, so to say, in quantity. Both interpretations harmonize with the observed fact that cases in which a female assumes male characters are much more frequent and much more definite than cases in which a male assumes femaleness.
Theory of Sexual Dimorphism.—Males and females may be alike, apart from their possession of male or female gonads, or may differ to almost any degree. It is plain, therefore, that although the presence and the maturity of the gonads may be, and probably are, the immediate stimulus to the appearance of the secondary differences, they cannot be the prime cause. Why, although equally potent sexually, do some males and females differ, others resemble one another? This is a question distinct from that of the primary determination of sex and the mechanism by which it is brought about. C. Darwin's theory of sexual selection remains the only comprehensive suggestion. Like his theory of the Origin of Species, it is not a theory of the origin of variations. He starts from the observed fact that variations occur and are transmitted; he supposes that by natural selection individuals favoured by suitable variations are preserved, and that in such a fashion the divergence which leads to the origin of species has come about; he also supposes that by sexual selection, or preferential mating, the differences between male and female have been brought about. “Courage, pugnacity, perseverance, strength and size of body, weapons of all kinds, musical organs, both vocal and instrumental, bright colours, stripes and marks, and ornamental appendages, have all been indirectly gained by the one sex or the other, through the influence of love and jealousy, through the appreciation of the beautiful in sound, colour or form, and through the exertion of a choice; and these powers of the mind manifestly depend on the development of the cerebral system” (Descent of Man, ii. p. 402). The characters to be accounted for are confined to one sex and are in closing relation with the breeding season and breeding habits. In those cases where they differ from the females, the males are the most active in courtship, and the best armed, and are rendered the most attractive in various ways. They fight with their rivals for the possession of the female, or display their attractions before her, and either by conquest or by being preferred have an advantage over less favoured males. Darwin was in some doubt as to how far it could be shown that such favoured individuals had a chance of leaving more progeny, except in cases where males were polygamous or much more numerous than females, but he suggested that on the whole the more vigorous female would be the the first to breed and to choose the more attractive males, or be captured by the stronger males. A. R. Wallace was unable to accept the theory of sexual selection except in the most limited way, and in particular laid great stress on the want of evidence, to which Darwin himself has called attention, that females prefer more highly ornamented males. He thought that natural selection was sufficient to explain sexual differences such as the possession of weapons, scents and call-notes. With regard to colour and pattern, he regarded these as natural outcrops of specialized structure, better displayed in more vigorous animals, and therefore likely to increase under natural selection. The inconspicuous patterns and dull colours of females he believed to depend on natural selection, and to be associated with the greater need for females to be inconspicuous whilst engaged in their duties to their young. More recent writers have shown that in a large number of cases brilliant colours and patterns are in themselves really protective (see Colours of Animals), so that the facts left to be explained by the theory of sexual selection are still further restricted.
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(P. C. M.)