The Evolution of Man — Volume 2

by Ernst Haeckel

This applies also to the mucous lining of the olfactory organ, the nose. However, the development of this organ is much more interesting. Although the nose seems superficially to be simple and single, it really consists, in man and all other Gnathostomes, of two completely separated halves, the right and left cavities. They are divided by a vertical partition, so that the right nostril leads into the right cavity alone and the left nostril into the left cavity. They open internally (and separately) by the posterior nasal apertures into the pharynx, so that we can get direct into the gullet through the nasal passages without touching the mouth. This is the way the air usually passes in respiration; the mouth being closed, it goes through the nose into the gullet, and through the larynx and bronchial tubes into the lungs. The nasal cavities are separated from the mouth by the horizontal bony palate, to which is attached behind (as a dependent process) the soft palate with the uvula. In the upper and hinder parts of the nasal cavities the olfactory nerve, the first pair of cerebral nerves, expands in the mucous coat which clothes them. The terminal branches of it spread partly over the septum (partition), partly on the side walls of the internal cavities, to which are attached the turbinated bones. These bones are much more developed in many of the higher mammals than in man, but there are three of them in all mammals. The sensation of smell arises by the passage of a current of air containing odorous matter over the mucous lining of the cavities, and stimulating the olfactory cells of the nerve-endings.

Man has all the features which distinguish the olfactory organ of the mammals from that of the lower Vertebrates. In all essential points the human nose entirely resembles that of the Catarrhine apes, some of which have quite a human external nose (compare the face of the long-nosed apes). However, the first structure of the olfactory organ in the human embryo gives no indication of the future ample proportions of our catarrhine nose. It has the form in which we find it permanently in the fishes—a couple of simple depressions in the skin at the outer surface of the head. We find these blind olfactory pits in all the fishes; sometimes they lie near the eyes, sometimes more forward at the point of the muzzle, sometimes lower down, near the mouth (Figure 2.249).

(FIGURE 2.311. Frontal section of the mouth and throat of a human embryo, neck half-inch long. "Invented" by Wilhelm His. The vertical section (in the frontal plane, from left to right) is so constructed that we see the nasal pits in the upper third of the figure and the eyes at the sides: in the middle third the primitive gullet with the gill-clefts (gill-arches in section); in the lower third the pectoral cavity with the bronchial tubes and the rudimentary lungs.)

This first rudimentary structure of the double nose is the same in all the Gnathostomes; it has no connection with the primitive mouth. But even in a section of the fishes a connection of this kind begins to make its appearance, a furrow in the surface of the skin running from each side of the nasal pit to the nearest corner of the mouth. This furrow, the nasal groove or furrow (Figure 2.305 r), is very important. In many of the sharks, such as the Scyllium, a special process of the frontal skin, the nasal fold or internal nasal process, is formed internally over the groove (n, n apostrophe). In contrast to this the outer edge of the furrow rises in an "external nasal process." As the two processes meet and coalesce over the nasal groove in the Dipneusts and Amphibia, it is converted into a canal, the nasal canal. Henceforth we can penetrate from the external pits through the nasal canals direct into the mouth, which has been formed quite independently. In the Dipneusts and the lower Amphibia the internal aperture of the nasal canals lies in front (behind the lips); in the higher Amphibia it is right behind. Finally, in the three higher classes of Vertebrates the primary mouth-cavity is divided by the formation of the horizontal palate-roof into two distinct cavities—the upper (secondary) nasal cavity and the lower (secondary) mouth-cavity. The nasal cavity in turn is divided by the construction of the vertical septum into two halves—right and left.

(FIGURE 2.312. Diagrammatic section of the mouth-nose cavity. While the palate-plates (p) divide the original mouth-cavity into the lower secondary mouth (m) and the upper nasal cavity, the latter in turn is divided by the vertical partition (e) into two halves (n, n). (From Gegenbaur.))

Comparative anatomy shows us to-day, in the series of the double-nosed Vertebrates, from the fishes up to man, all the different stages in the development of the nose, which the advanced olfactory organ of the higher mammals has passed through at various periods in the course of its phylogeny. It first appears in the embryo of man and the higher Vertebrates, in which the double fish-nose persists throughout life. At an early stage, before there is any trace of the characteristic human face, a pair of small pits are formed in the head over the original mouth-cavity; these were first discovered by Baer, and rightly called the "olfactory pits" (Figures 2.306 n and 2.307 n). These primitive nasal pits are quite separate from the rudimentary mouth, which also originates as a pit-like depression in the skin, in front of the blind fore end of the gut. Both the pair of nasal pits and the single mouth-pit (Figure 2.310 m) are clothed with the horny plate. The original separation of the former from the latter is, however, presently abolished, a process forming above the mouth-pit—the "frontal process" (Figure 2.309 st). Its outer edge rises to the right and left in the shape of two lateral processes; these are the inner nasal processes or folds (in). Opposite to these a parallel ridge is formed on either side between the eye and the nasal pit; these are the outer nasal processes (an). Thus between the inner and outer nasal processes a groove-like depression is formed on either side, which leads from the nasal pit towards the mouth-pit (m); this groove is, as the reader will guess, the same nasal furrow or groove that we have already seen in the shark (Figure 2.305 r). As the parallel edges of the inner and outer nasal processes bend towards each other and join above the nasal groove, this is converted into a tube, the primitive nasal canal. Hence the nose of man and all the other Amniotes consists at this embryonic stage of a couple of narrow tubes, the nasal canals, which lead from the outer surface of the forehead into the rudimentary mouth. This transitory condition resembles that in which we find the nose permanently in the Dipneusts and Amphibia.

A cone-shaped structure, which grows from below towards the lower ends of the two nasal processes and joins with them, plays an important part in the conversion of the open nasal groove into the closed canal. This is the upper-jaw process (Figures 2.306 to 2.310 o). Below the mouth-pit are the gill-arches, which are separated by the gill-clefts. The first of these gill-arches, and the most important for our purpose, which we may call the maxillary (jaw) arch, forms the skeleton of the jaws. Above at the basis a small process grows out of this first gill-arch; this is the upper-jaw process. The first gill-arch itself develops a cartilage at one of its inner sides, the "Meckel cartilage" (named after its discoverer), on the outer surface of which the lower jaw is formed (Figures 2.306 to 2.310 u). The upper-jaw process forms the chief part of the skeleton of that jaw, the palate bone, and the pterygoid bone. On its outer side is afterwards formed the upper-jaw bone, in the narrower sense, while the middle part of the skeleton of the upper jaw, the intermaxillary, develops from the foremost part of the frontal process.

The two upper-jaw processes are of great importance in the further development of the face. From them is formed, growing into the primitive mouth-cavity, the important horizontal partition (the palate) that divides the former into two distinct cavities. The upper cavity, into which the nasal canals open, now develops into the nasal cavity, the air-passage and the organ of smell. The lower cavity forms the permanent secondary mouth (Figure 2.312 m), the food-passage and the organ of taste. Both the upper and lower cavities open behind into the gullet (pharynx). The hard palate that separates them is formed by the joining of two lateral halves, the horizontal plates of the two upper-jaw processes, or the palate-plates (p). When these do not, sometimes, completely join in the middle, a longitudinal cleft remains, through which we can penetrate from the mouth straight into the nasal cavity. This is the malformation known as "wolf's throat." "Hare-lip" is the lesser form of the same defect. At the same time as the horizontal partition of the hard palate a vertical partition is formed by which the single nasal cavity is divided into two sections—a right and left half (Figure 2.312 n, n).

(FIGURES 2.313 AND 2.314. Upper part of the body of a human embryo, two-thirds of an inch long, of the sixth week; Figure 2.313 from the left, Figure 2.314 from the front. The origin of the nose and the upper lip from two lateral and originally separate halves can be clearly seen. Nose and upper lip are large in proportion to the rest of the face, and especially to the lower lip. (From Kollmann.))

The double nose has now acquired the characteristic form that man shares with the other mammals. Its further development is easy to follow; it consists of the formation of the inner and outer processes of the walls of the two cavities. The external nose is not formed until long after all these essential parts of the internal organ of smell. The first traces of it in the human embryo are found about the middle of the second month (Figures 2.313 to 2.316). As can be seen in any human embryo during the first month, there is at first no trace of the external nose. It only develops afterwards from the foremost nasal part of the primitive skull, growing forwards from behind. The characteristic human nose is formed very late. Much stress is at times laid on this organ as an exclusive privilege of man. But there are apes that have similar noses, such as the long-nosed ape.