Chapter VII: The Forms of Tissues or Cell-Aggregates (2)

While the formation of an hexagonal pattern on the basis of ready-formed and symmetrically arranged material units is a very common, and indeed the general way, it does not follow that there are not others by which such a pattern can be obtained. For instance, if we take a little triangular dish of mercury and set it vibrating (either by help of a tuning-fork, or by simply tapping on the sides) we shall have a series of little waves or ripples starting inwards from each of the three faces; and the intercrossing, or interference of these three sets of waves produces crests and hollows, and intermediate points of no disturbance, _whose loci are seen_ as a beautiful pattern of minute hexagons. It is possible that the very minute and astonishingly regular pattern of hexagons which we see, for instance, on the surface of many diatoms, may be a phenomenon of this order[363]. The same may be the case also in Arcella, where an apparently hexagonal pattern is found not to consist of simple hexagons, but of “straight lines in three sets of parallels, the lines of each set making an angle of sixty degrees with those of the other two sets[364].” We must also bear in mind, in the case of the minuter forms, the large possibilities of optical illusion. For instance, in one of Abbe’s “diffraction-plates,” a pattern of dots, set at equal interspaces, is reproduced on a very minute scale by photography; but under certain conditions of microscopic illumination and focussing, these isolated dots appear as a pattern of hexagons.

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A symmetrical arrangement of hexagons, such as we have just been studying, suggests various simple geometrical corollaries, of which the following may perhaps be a useful one.

We may sometimes desire to estimate the number of hexagonal areas or facets in some structure where these are numerous, such for instance as the {324} cornea of an insect’s eye, or in the minute pattern of hexagons on many diatoms. An approximate enumeration is easily made as follows.

For the area of a hexagon (if we call δ the short diameter, that namely which bisects two of the opposite sides) is δ^2 × (√3)/2, the area of a circle being _d_^2 ⋅ π/4. Then, if the diameter (_d_) of a circular area include _n_ hexagons, the area of that circle equals (_n_ ⋅ δ)^2 × π/4. And, dividing this number by the area of a single hexagon, we obtain for the number of areas in the circle, each equal to a hexagonal facet, the expression _n_^2 × π/4 × 2/(√3) = 0·907_n_^2, or (9/10) ⋅ _n_^2, nearly.

This calculation deals, not only with the complete facets, but with the areas of the broken hexagons at the periphery of the circle. If we neglect these latter, and consider our whole field as consisting of successive rings of hexagons about a central one, we may obtain a still simpler rule[365]. For obviously, around our central hexagon there stands a zone of six, and around these a zone of twelve, and around these a zone of eighteen, and so on. And the total number, excluding the central hexagon, is accordingly:

For one zone 6 = 2 × 3 = 3 × 1 × 2, For two zones 18 = 3 × 6 = 3 × 2 × 3, For three zones 36 = 4 × 9 = 3 × 3 × 4, For four zones 60 = 5 × 12 = 3 × 4 × 5, For five zones 90 = 6 x 15 = 3 × 5 × 6,

and so forth. If _N_ be the number of zones, and if we add one to the above numbers for the odd central hexagon, the rule evidently is, that the total number, _H_, = 3_N_(_N_ + 1) + 1. Thus, if in a preparation of a fly’s cornea, I can count twenty-five facets in a line from a central one, the total number in the entire circular field is (3 × 25 × 26) + 1 = 1951[366].

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The same principles which account for the development of hexagonal symmetry hold true, as a matter of course, not only of individual _cells_ (in the biological sense), but of any close-packed bodies of uniform size and originally circular outline; and the hexagonal pattern is therefore of very common occurrence, under widely different circumstances. The curious reader may consult Sir Thomas Browne’s quaint and beautiful account, in the _Garden of Cyrus_, of hexagonal (and also of quincuncial) symmetry in plants and animals, which “doth neatly declare how nature Geometrizeth, and observeth order in all things.” {325}

We have many varied examples of this principle among corals, wherever the polypes are in close juxtaposition, with neither empty space nor accumulations of matrix between their adjacent walls. _Favosites gothlandica_, for instance, furnishes us with an excellent example. In the great genus Lithostrotion we have some species that are “massive” and others that are “fasciculate”; in other words in some the long cylindrical corallites are in close contact with one another, and in others they are separate and loosely bundled (Fig. 127). Accordingly in the former the corallites are

squeezed into hexagonal prisms, while in the latter they retain their cylindrical form. Where the polypes are comparatively few, and so have room to spread, the mutual pressure ceases to work or only tends to push them asunder, letting them remain circular in outline (e.g. Thecosmilia). Where they vary gradually in size, as for instance in _Cyathophyllum hexagonum_, they are more or less hexagonal but are not regular hexagons; and where there is greater and more irregular variation in size, the cells will be _on the average_ hexagonal, but some will have fewer and some more sides than six, as in the annexed figure of Arachnophyllum (Fig. 129). {326} Where larger and smaller cells, corresponding to two different kinds of zooids, are mixed together, we may get various results. If the larger cells are numerous enough to be more or less in contact with one another (e.g. various Monticuliporae) they will be irregular hexagons, while the smaller cells between them will be crushed into all manner of irregular angular forms. If on the other hand the large cells are comparatively few and are large and strong-walled compared with their smaller neighbours, then the latter alone will be squeezed into hexagons, while the larger ones will tend to retain their circular outline undisturbed (e.g. Heliopora, Heliolites, etc.).

When, as happens in certain corals, the peripheral walls or “thecae” of the individual polypes remain undeveloped but the radiating septa are formed and calcified, then we obtain new and beautiful mathematical configurations (Fig. 131). For the radiating septa are no longer confined to the circular or hexagonal bounds of a polypite, but tend to meet and become confluent with their neighbours on every side; and, tending to assume positions of equilibrium, or of minimal area, under the restraints to which they are subject, they fall into congruent curves; and these correspond, in a striking manner, to the lines of force running, in a common field of force, between a number of secondary centres. Similar patterns may be produced in various ways, by the play of osmotic or magnetic forces; and a particular and very curious case is to be found in those complicated forms of nuclear division {327} known as triasters, polyasters, etc., whose relation to a field of force Hartog has explained[367]. It is obvious that, in our corals, these curving septa are all orthogonal to the non-existent hexagonal boundaries. As the phenomenon is wholly due to the imperfect development or non-existence of a thecal wall, it is not surprising that we find identical configurations among various corals, or families of corals, not otherwise related to one another; we find the same or very similar patterns displayed, for instance, in Synhelia (_Oculinidae_), in Phillipsastraea (_Rugosa_), in Thamnastraea (_Fungida_), and in many more.

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The most famous of all hexagonal conformations and perhaps the most beautiful is that of the bee’s cell. Here we have, as in our last examples, a series of equal cylinders, compressed by symmetrical forces into regular hexagonal prisms. But in this case we have two rows of such cylinders, set opposite to one another, end to end; and we have accordingly to consider also the conformation of their ends. We may suppose our original cylindrical cells to have spherical ends, which is their normal and symmetrical mode of termination; and, for closest packing, it is obvious that the end of any one cylinder will touch, and fit in between, the ends of three cylinders in the opposite row. It is just as when we pile round-shot in a heap; each sphere that we {328} set down fits into its nest between three others, and the four form a regular tetrahedral arrangement. Just as it was obvious, then, that by mutual pressure from the six _laterally_ adjacent cells, any one cell would be squeezed into a hexagonal prism, so is it also obvious that, by mutual pressure against the three _terminal_ neighbours, the end of any one cell will be compressed into a solid trihedral angle whose edges will meet, as in the analogous case already described of a system of soap-bubbles, at a plane angle of 109° and so many minutes and seconds. What we have to comprehend, then, is how the _six_ sides of the cell are to be combined with its _three_ terminal facets. This is done by bevelling off three alternate angles of the prism, in a uniform manner, until we have tapered the prism to a point; and by so doing, we evidently produce three _rhombic_ surfaces, each of which is double of the triangle formed by joining the apex to the three untouched angles of the prism. If we experiment, not with cylinders, but with spheres, if for instance we pile together a mass of bread-pills (or pills of plasticine), and then submit the whole to a uniform pressure, it is obvious that each ball (like the seeds in a pomegranate, as Kepler said), will be in contact with _twelve_ others,—six in its own plane, three below and three above, and in compression it will therefore develop twelve plane surfaces. It will in short repeat, above and below, the conditions to which the bee’s cell is subject at one end only; and, since the sphere is symmetrically situated towards its neighbours on all sides, it follows that the twelve plane sides to which its surface has been reduced will be all similar, equal and similarly situated. Moreover, since we have produced this result by squeezing our original spheres close together, it is evident that the bodies so formed completely fill space. The regular solid which fulfils all these conditions is the _rhombic dodecahedron_. The bee’s cell, then, is this figure incompletely formed: it is a hexagonal prism with one open or unfinished end, and one trihedral apex of a rhombic dodecahedron.

The geometrical form of the bee’s cell must have attracted the attention and excited the admiration of mathematicians from time immemorial. Pappus the Alexandrine has left us (in the introduction to the Fifth Book of his _Collections_) an account of its hexagonal plan, and he drew from its mathematical symmetry the {329} conclusion that the bees were endowed with reason: “There being, then, three figures which of themselves can fill up the space round a point, viz. the triangle, the square and the hexagon, the bees have wisely selected for their structure that which contains most angles, suspecting indeed that it could hold more honey than either of the other two.” Erasmus Bartholinus was apparently the first to suggest that this hypothesis was not warranted, and that the hexagonal form was no more than the necessary result of equal pressures, each bee striving to make its own little circle as large as possible.

The investigation of the ends of the cell was a more difficult matter, and came later, than that of its sides. In general terms this arrangement was doubtless often studied and described: as for instance, in the _Garden of Cyrus_: “And the Combes themselves so regularly contrived that their mutual intersections make three Lozenges at the bottom of every Cell; which severally regarded make three Rows of neat Rhomboidall Figures, connected at the angles, and so continue three several chains throughout the whole comb.” But Maraldi[368] (Cassini’s nephew) was the first to measure the terminal solid angle or determine the form of the rhombs in the pyramidal ending of the cell. He tells us that the angles of the rhomb are 110° and 70°: “Chaque base d’alvéole est formée par trois rhombes presque toujours égaux et semblables, qui, suivant les mesures que nous avons prises, ont les deux angles obtus chacun de 110 degrés, et par conséquent les deux aigus chacun de 70°.” He also stated that the angles of the trapeziums which form the sides of the body of the cell were identical angles, of 110° and 70°; but in the same paper he speaks of the angles as being, respectively, 109° 28′ and 70° 32′. Here a singular confusion at once arose, and has been perpetuated in the books[369]. “Unfortunately Réaumur chose to look upon this second determination of Maraldi’s as being, as well as the first, a direct result of measurement, whereas it is in reality theoretical. He speaks of it as Maraldi’s more precise measurement, and this error has been repeated in spite of its absurdity to the present day; nobody {330} appears to have thought of the impossibility of measuring such a thing as the end of a bee’s cell to the nearest minute.” At any rate, it now occurred to Réaumur (as curiously enough, it had not done to Maraldi) that, just as the closely packed hexagons gave the minimal extent of boundary in a plane, so the actual solid figure, as determined by Maraldi, might be that which, for a given solid content, gives the minimum of surface: or which, in other words, would hold the most honey for the least wax. He set this problem before Koenig, and the geometer confirmed his conjecture, the result of his calculations agreeing within two minutes (109° 26′ and 70° 34′) with Maraldi’s determination. But again, Maclaurin[370] and Lhuilier[371], by different methods, obtained a result identical with Maraldi’s; and were able to shew that the discrepancy of 2′ was due to an error in Koenig’s calculation (of tan θ = √2),—that is to say to the imperfection of his logarithmic tables,—not (as the books say[372]) “to a mistake on the part of the Bee.” “Not to a mistake on the part of Maraldi” is, of course, all that we are entitled to say.

The theorem may be proved as follows:

_ABCDEF_, _abcdef_, is a right prism upon a regular hexagonal base. The corners _BDF_ are cut off by planes through the lines _AC_, _CE_, _EA_, meeting in a point _V_ on the axis _VN_ of the prism, and intersecting _Bb_, _Dd_, _Ff_, at _X_, _Y_, _Z_. It is evident that the volume of the figure thus formed is the same as that of the original prism with hexagonal ends. For, if the axis cut the hexagon _ABCDEF_ in _N_, the volumes _ACVN_, _ACBX_ are equal. {331}

It is required to find the inclination of the faces forming the trihedral angle at _V_ to the axis, such that the surface of the figure may be a minimum.

Let the angle _NVX_, which is half the solid angle of the prism, = θ; the side of the hexagon, as _AB_, = _a_; and the height, as _Aa_, = _h_.

Then, _AC_ = 2_a_ cos 30° = _a_√3.

And _VX_ = _a_/sin θ (from inspection of the triangle _LXB_)

Therefore the area of the rhombus _VAXC_ = (_a_^2 √3)/(2 sin θ).

And the area of _AabX_ = (_a_/2)(2_h_ − ½_VX_ cos θ)

= (_a_/2)(2_h_ − _a_/2 ⋅ cot θ).

Therefore the total area of the figure

= hexagon _abcdef_ + 3_a_(2_h_ − (_a_/2) cot θ)
+ 3((_a_^2 √3)/(2 sin θ)).

Therefore _d_(Area)/_d_θ = (3_a_^2/2)((1/sin^2 θ)
− (√3 cos θ)/(sin^2 θ)).

But this expression vanishes, that is to say, _d_(Area)/_d_θ = 0, when cos θ = 1/√3, that is when θ = 54° 44′ 8″ = ½(109° 28′ 16″).

This then is the condition under which the total area of the figure has its minimal value.

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That the beautiful regularity of the bee’s architecture is due to some automatic play of the physical forces, and that it were fantastic to assume (with Pappus and Réaumur) that the bee intentionally seeks for a method of economising wax, is certain, but the precise manner of this automatic action is not so clear. When the hive-bee builds a solitary cell, or a small cluster of cells, as it does for those eggs which are to develop into queens, it makes but a rude production. The queen-cells are lumps of coarse wax hollowed out and roughly bitten into shape, bearing the marks of the bee’s jaws, like the marks of a blunt adze on a rough-hewn log. Omitting the simplest of all cases, when (as among some humble-bees) the old cocoons are used to hold honey, the cells built by the “solitary” wasps and bees are of various kinds. They may be formed by partitioning off little chambers in a hollow stem; {332} they may be rounded or oval capsules, often very neatly constructed, out of mud, or vegetable _fibre_ or little stones, agglutinated together with a salivary glue; but they shew, except for their rounded or tubular form, no mathematical symmetry. The social wasps and many bees build, usually out of vegetable matter chewed into a paste with saliva, very beautiful nests of “combs”; and the close-set papery cells which constitute these combs are just as regularly hexagonal as are the waxen cells of the hive-bee. But in these cases (or nearly all of them) the cells are in a single row; their sides are regularly hexagonal, but their ends, from the want of opponent forces, remain simply spherical. In _Melipona domestica_ (of which Darwin epitomises Pierre Huber’s description) “the large waxen honey-cells are nearly spherical, nearly equal in size, and are aggregated into an irregular mass.” But the spherical form is only seen on the outside of the mass; for inwardly each cell is flattened into “two, three or more flat surfaces, according as the cell adjoins two, three or more other cells. When one cell rests on three other cells, which from the spheres being nearly of the same size is very frequently and necessarily the case, the three flat surfaces are united into a pyramid; and this pyramid, as Huber has remarked, is manifestly a gross imitation of the three-sided pyramidal base of the cell of the hive-bee[373].” The question is, to what particular force are we to ascribe the plane surfaces and definite angles which define the sides of the cell in all these cases, and the ends of the cell in cases where one row meets and opposes another. We have seen that Bartholin suggested, and it is still commonly believed, that this result is due to simple physical pressure, each bee enlarging as much as it can the cell which it is a-building, and nudging its wall outwards till it fills every intervening gap and presses hard against the similar efforts of its neighbour in the cell next door[374]. But it is very doubtful {333} whether such physical or mechanical pressure, more or less intermittently exercised, could produce the all but perfectly smooth, plane surfaces and the all but perfectly definite and constant angles which characterise the cell, whether it be constructed of wax or papery pulp. It seems more likely that we have to do with a true surface-tension effect; in other words, that the walls assume their configuration when in a semi-fluid state, while the papery pulp is still liquid, or while the wax is warm under the high temperature of the crowded hive[375]. Under these circumstances, the direct efforts of the wasp or bee may be supposed to be limited to the making of a tubular cell, as thin as the nature of the material permits, and packing these little cells as close as possible together. It is then easily conceivable that the symmetrical tensions of the adjacent films (though somewhat retarded by viscosity) should suffice to bring the whole system into equilibrium, that is to say into the precise configuration which the comb actually presents. In short, the Maraldi pyramids which terminate the bee’s cell are precisely identical with the facets of a rhombic dodecahedron, such as we have assumed to constitute (and which doubtless under certain conditions do constitute) the surfaces of contact in the interior of a mass of soap-bubbles or of uniform parenchymatous cells; and there is every reason to believe that the physical explanation is identical, and not merely mathematically analogous.

The remarkable passage in which Buffon discusses the bee’s cell and the hexagonal configuration in general is of such historical importance, and tallies so closely with the whole trend of our enquiry, that I will quote it in full: “Dirai-je encore un mot; ces cellules des abeilles, tant vantées, tant admirées, me fournissent une preuve de plus contre l’enthousiasme et l’admiration; cette figure, toute géométrique et toute régulière qu’elle nous paraît, et qu’elle est en effet dans la spéculation, n’est ici qu’un résultat mécanique et assez imparfait qui se trouve souvent dans la nature, {334} et que l’on remarque même dans les productions les plus brutes; les cristaux et plusieurs autres pierres, quelques sels, etc., prennent constamment cette figure dans leur formation. Qu’on observe les petites écailles de la peau d’une roussette, on verra qu’elles sont hexagones, parce que chaque écaille croissant en même temps se fait obstacle, et tend à occuper le plus d’espace qu’il est possible dans un espace donné: on voit ces mêmes hexagones dans le second estomac des animaux ruminans, on les trouve dans les graines, dans leurs capsules, dans certaines fleurs, etc. Qu’on remplisse un vaisseau de pois, ou plûtot de quelque autre graine cylindrique, et qu’on le ferme exactement après y avoir versé autant d’eau que les intervalles qui restent entre ces graines peuvent en recevoir; qu’on fasse bouillir cette eau, tous ces cylindres deviendront de colonnes à six pans[376]. On y voit clairement la raison, qui est purement mécanique; chaque graine, dont la figure est cylindrique, tend par son renflement à occuper le plus d’espace possible dans un espace donné, elles deviennent donc toutes nécessairement hexagones par la compression réciproque. Chaque abeille cherche à occuper de même le plus d’espace possible dans un espace donné, il est donc nécessaire aussi, puisque le corps des abeilles est cylindrique, que leurs cellules sont hexagones,—par la même raison des obstacles réciproques. On donne plus d’esprit aux mouches dont les ouvrages sont les plus réguliers; les abeilles sont, dit-on, plus ingénieuses que les guêpes, que les frélons, etc., qui savent aussi l’architecture, mais dont les constructions sont plus grossières et plus irrégulières que celles des abeilles: on ne veut pas voir, ou l’on ne se doute pas que cette régularité, plus ou moins grande, dépend uniquement du nombre et de la figure, et nullement de l’intelligence de ces petites bêtes; plus elles sont nombreuses, plus il y a des forces qui agissent également et s’opposent de même, plus il y a par conséquent de contrainte mécanique, de régularité forcée, et de perfection apparente dans leurs productions[377].” {335}

A very beautiful hexagonal symmetry, as seen in section, or dodecahedral, as viewed in the solid, is presented by the cells which form the pith of certain rushes (e.g. _Juncus effusus_), and somewhat less diagrammatically by those which make the pith of the banana. These cells are stellate in form, and the tissue presents in section the appearance of a network of six-rayed stars (Fig. 133, _c_), linked together by the tips of the rays, and separated by symmetrical, air-filled, intercellular spaces. In thick sections, the solid twelve-rayed stars may be very beautifully seen under the binocular microscope.

What has happened here is not difficult to understand. Imagine, as before, a system of equal spheres all in contact, each one therefore touching six others in an equatorial plane; and let the cells be not only in contact, but become attached at the points of contact. Then instead of each cell expanding, so as to encroach on and fill up the intercellular spaces, let each cell tend to contract or shrivel up, by the withdrawal of fluid from its interior. The {336} result will obviously be that the intercellular spaces will increase; the six equatorial attachments of each cell (Fig. 133, _a_) (or its twelve attachments in all, to adjacent cells) will remain fixed, and the portions of cell-wall between these points of attachment will be withdrawn in a symmetrical fashion (_b_) towards the centre. As the final result (_c_) we shall have a “dodecahedral star” or star-polygon, which appears in section as a six-rayed figure. It is obviously necessary that the pith-cells should not only be attached to one another, but that the outermost layer should be firmly attached to a boundary wall, so as to preserve the symmetry of the system. What actually occurs in the rush is tantamount to this, but not absolutely identical. Here it is not so much the pith-cells which tend to shrivel within a boundary of constant size, but rather the boundary wall (that is, the peripheral ring of woody and other tissues) which continues to expand after the pith-cells which it encloses have ceased to grow or to multiply. The twelve points of attachment on the spherical surface of each little pith-cell are uniformly drawn asunder; but the content, or volume, of the cell does not increase correspondingly; and the remaining portions of the surface, accordingly, shrink inwards and gradually constitute the complicated surface of a twelve-pointed star, which is still a symmetrical figure and is still also a surface of minimal area under the new conditions.

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A few years after the publication of Plateau’s book, Lord Kelvin shewed, in a short but very beautiful paper[378], that we must not hastily assume from such arguments as the foregoing, that a close-packed assemblage of rhombic dodecahedra will be the true and general solution of the problem of dividing space with a minimum partitional area, or will be present in a cellular liquid “foam,” in which it is manifest that the problem is actually and automatically solved. The general mathematical solution of the problem (as we have already indicated) is, that every interface or partition-wall must have constant curvature throughout; that where such partitions meet in an edge, they must intersect at angles such that equal forces, in planes perpendicular to the line {337} of intersection, shall balance; and finally, that no more than three such interfaces may meet in a line or edge, whence it follows that the angle of intersection of the film-surfaces must be exactly 120°. An assemblage of equal and similar rhombic dodecahedra goes far to meet the case: it completely fills up space; all its surfaces or interfaces are planes, that is to say, surfaces of constant curvature throughout; and these surfaces all meet together at angles of 120°. Nevertheless, the proof that our rhombic dodecahedron (such as we find exemplified in the bee’s cell) is a surface of minimal area, is not a comprehensive proof; it is limited to certain conditions, and practically amounts to no more than this, that of the regular solids, with all sides plane and similar, this one has the least surface for its solid content.

The rhombic dodecahedron has six tetrahedral angles, and eight trihedral angles; and it is obvious, on consideration, that at each of the former six dodecahedra meet in a point, and that, where the four tetrahedral facets of each coalesce with their neighbours, we have twelve plane films, or interfaces, meeting in a point. In a precisely similar fashion, we may imagine twelve plane films, drawn inwards from the twelve edges of a cube, to meet at a point in the centre of the cube. But, as Plateau discovered[379], when we dip a cubical wire skeleton into soap-solution and take it out again, the twelve films which are thus generated do _not_ meet in a point, but are grouped around a small central, plane, quadrilateral film (Fig. 134). In other words, twelve plane films, meeting in a point, are _essentially unstable_. If we blow upon our artificial film-system, the little quadrilateral alters its place, setting itself parallel now to one and now to another of the paired faces of the cube; but we never get rid of it. Moreover, the size and shape of the quadrilateral, as of all the other films in the system, are perfectly definite. Of the twelve films (which we had {338} expected to find all plane and all similar) four are plane isosceles triangles, and eight are slightly curved quadrilateral figures. The former have two curved sides, meeting at an angle of 109° 28′, and their apices coincide with the corners of the central quadrilateral, whose sides are also curved, and also meet at this identical angle;—which (as we observe) is likewise an angle which we have been dealing with in the simpler case of the bee’s cell, and indeed in all the regular solids of which we have yet treated.

By completing the assemblage of polyhedra of which Plateau’s skeleton-cube gives a part, Lord Kelvin shewed that we should obtain a set of equal and similar fourteen-sided figures, or “tetrakaidecahedra”; and that by means of an assemblage of these figures space is homogeneously partitioned—that is to say, into equal, similar and similarly situated cells—with an economy of surface in relation to area even greater than in an assemblage of rhombic dodecahedra.

In the most generalised case, the tetrakaidecahedron is bounded by three pairs of equal and parallel quadrilateral faces, and four pairs of equal and parallel hexagonal faces, neither the quadrilaterals nor the hexagons being necessarily plane. In a certain particular case, the quadrilaterals are plane surfaces, but the hexagons slightly curved “anticlastic” surfaces; and these latter have at every point equal and opposite curvatures, and are surfaces of minimal curvature for a boundary of six curved edges. The figure has the remarkable property that, like the plane rhombic dodecahedron, it so partitions space that three faces meeting in an edge do so everywhere at equal angles of 120° [380].

We may take it as certain that, in a system of _perfectly_ fluid films, like the interior of a mass of soap-bubbles, where the films are perfectly free to glide or to rotate over one another, the mass is actually divided into cells of this remarkable conformation. {339} And it is quite possible, also, that in the cells of a vegetable parenchyma, by carefully macerating them apart, the same conformation may yet be demonstrated under suitable conditions; that is to say when the whole tissue is highly symmetrical, and the individual cells are as nearly as possible equal in size. But in an ordinary microscopic _section_, it would seem practically impossible to distinguish the fourteen-sided figure from the twelve-sided. Moreover, if we have anything whatsoever interposed so as to prevent our twelve films meeting in a point, and (so to speak) to take the place of our little central quadrilateral,—if we have, for instance, a tiny bead or droplet in the centre of our artificial system, or even a little thickening, or “bourrelet” as Plateau called it, of the cell-wall, then it is no longer necessary that the tetrakaidecahedron should be formed. Accordingly, it is very probably the case that, in the parenchymatous tissue, under the actual conditions of restraint and of very imperfect fluidity, it is after all the rhombic dodecahedral configuration which, even under perfectly symmetrical conditions, is generally assumed.

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It follows from all that we have said, that the problems connected with the conformation of cells, and with the manner in which a given space is partitioned by them, soon become exceedingly complex. And while this is so even when all our cells are equal and symmetrically placed, it becomes vastly more so when cells varying even slightly in size, in hardness, rigidity or other qualities, are packed together. The mathematics of the case very soon become too hard for us; but in its essence, the phenomenon remains the same. We have little reason to doubt, and no just cause to disbelieve, that the whole configuration, for instance of an egg in the advanced stages of segmentation, is accurately determined by simple physical laws, just as much as in the early stages of two or four cells, during which early stages we are able to recognise and demonstrate the forces and their resultant effects. But when mathematical investigation has become too difficult, it often happens that physical experiment can reproduce for us the phenomena which Nature exhibits to us, and which we are striving to comprehend. For instance, in an admirable research, M. Robert shewed, some years ago, not only that the early segmentation of {340} the egg of _Trochus_ (a marine univalve mollusc) proceeded in accordance with the laws of surface tension, but he also succeeded in imitating by means of soap-bubbles, several stages, one after another, of the developing egg.

M. Robert carried his experiments as far as the stage of sixteen cells, or bubbles. It is not easy to carry the artificial system quite so far, but in the earlier stages the experiment is easy; we have merely to blow our bubbles in a little dish, adding one to another, and adjusting their sizes to produce a symmetrical system. One of the simplest and prettiest parts of his investigation concerned the “polar furrow” of which we have spoken on p. 310. On blowing four little contiguous bubbles he found (as we may all find with the greatest ease) that they form a symmetrical system, two in contact with one another by a laminar film, and two, which are elevated a little above the others, and which are separated by the length of the aforesaid lamina. The bubbles are thus in contact three by three, their partition-walls making with one another equal angles of 120°. The upper and lower edges of the intermediate lamina (the lower one visible through the transparent system) constitute the two polar furrows of the embryologist (Fig. 135, 1–3). The lamina itself is plane when the system is symmetrical, but it responds by a corresponding curvature to the least inequality of the bubbles on either side. In the experiment, the upper polar furrow is usually a little shorter than the lower, but parallel to it; that is to say, the lamina is of trapezoidal form: this lack of perfect symmetry being due (in the experimental case) to the lower portion of the bubbles being somewhat drawn asunder by the tension of their attachments to the sides of the dish (Fig. 135, 4). A similar phenomenon is usually found in Trochus, according to Robert, and many other observers have likewise found the upper furrow to be shorter than the one below. In the various species of the genus Crepidula, Conklin asserts that the two furrows are equal in _C. convexa_, that the upper one is the shorter in _C. fornicata_, and that the upper one all but disappears in _C. plana_; but we may well be permitted to doubt, without the evidence of very special investigations, whether these slight physical differences are actually characteristic of, and constant in, particular allied _species_. {341} Returning to the experimental case, Robert found that by withdrawing a little air from, and so diminishing the bulk of the two terminal bubbles (i.e. those at the ends of the intermediate lamina), the upper polar furrow was caused to elongate, till it became equal in length to the lower; and by continuing the process it became the longer in its turn. These two conditions have again been described by investigators as characteristic of this embryo or that; for instance in Unio, Lillie has described the two furrows as gradually altering their respective lengths[381]; and Wilson (as Lillie remarks) had already pointed out that “the reduction of the apical cross-furrow, as compared with that at the vegetative pole {342} in molluscs and annelids ‘stands in obvious relation to the different size of the cells produced at the two poles[382].’ ”

When the two lateral bubbles are gradually reduced in size, or the two terminal ones enlarged, the upper furrow becomes shorter and shorter; and at the moment when it is about to vanish, a new furrow makes its instantaneous appearance in a direction perpendicular to the old one; but the inferior furrow, constrained by its attachment to the base, remains unchanged, and accordingly our two polar furrows, which were formerly parallel, are now at right angles to one another. Instead of a single plane quadrilateral partition, we have now two triangular ones, meeting in the middle of the system by their apices, and lying in planes at right angles to one another (Fig. 135, 5–7)[383]. Two such polar furrows, equal in length and arranged in a cross, have again been frequently described by the embryologists. Robert himself found this condition in Trochus, as an occasional or exceptional occurrence: it has been described as normal in Asterina by Ludwig, in Branchipus by Spangenberg, and in Podocoryne and Hydractinia by Bunting. It is evident that it represents a state of unstable equilibrium, only to be maintained under certain conditions of restraint within the system.

So, by slight and delicate modifications in the relative size of the cells, we may pass through all the possible arrangements of the median partition, and of the “furrows” which correspond to its upper and lower edges; and every one of these arrangements has been frequently observed in the four-celled stage of various embryos. As the phases pass one into the other, they are accompanied by changes in the curvature of the partition, which in like manner correspond precisely to phenomena which the embryologists have witnessed and described. And all these configurations belong to that large class of phenomena whose distribution among embryos, or among organisms in general, bears no relation to the boundaries of zoological classification; through molluscs, worms, {343} coelenterates, vertebrates and what not, we meet with now one and now another, in a medley which defies classification. They are not “vital phenomena,” or “functions” of the organism, or special characteristics of this or that organism, but purely physical phenomena. The kindred but more complicated phenomena which correspond to the polar furrow when a larger number of cells than four are associated together, we shall deal with in the next chapter.

Having shewn that the capillary phenomena are patent and unmistakable during the earlier stages of embryonic development, but soon become more obscure and incapable of experimental reproduction in the later stages, when the cells have increased in number, various writers including Robert himself have been inclined to argue that the physical phenomena die away, and are overpowered and cancelled by agencies of a very different order. Here we pass into a region where direct observation and experiment are not at hand to guide us, and where a man’s trend of thought, and way of judging the whole evidence in the case, must shape his philosophy. We must remember that, even in a froth of soap-bubbles, we can apply an exact analysis only to the simplest cases and conditions of the phenomenon; we cannot describe, but can only imagine, the forces which in such a froth control the respective sizes, positions and curvatures of the innumerable bubbles and films of which it consists; but our knowledge is enough to leave us assured that what we have learned by investigation of the simplest cases includes the principles which determine the most complex. In the case of the growing embryo we know from the beginning that surface tension is only one of the physical forces at work; and that other forces, including those displayed within the interior of each living cell, play their part in the determination of the system. But we have no evidence whatsoever that at this point, or that point, or at any, the dominion of the physical forces over the material system gives place to a new condition where agencies at present unknown to the physicist impose themselves on the living matter, and become responsible for the conformation of its material fabric.

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Before we leave for the present the subject of the segmenting {344} egg, we must take brief note of two associated problems: viz. (1) the formation and enlargement of the segmentation cavity, or central interspace around which the cells tend to group themselves in a single layer, and (2) the formation of the gastrula, that is to say (in a typical case) the conversion “by invagination,” of the one-layered ball into a two-layered cup. Neither problem is free from difficulty, and all we can do meanwhile is to state them in general terms, introducing some more or less plausible assumptions.

The former problem is comparatively easy, as regards the tendency of a segmentation cavity to _enlarge_, when once it has been established. We may then assume that subdivision of the cells is due to the appearance of a new-formed septum within each cell, that this septum has a tendency to shrink under surface tension, and that these changes will be accompanied on the whole by a diminution of surface energy in the system. This being so, it may be shewn that the volume of the divided cells must be less than it was prior to division, or in other words that part of their contents must exude during the process of segmentation[384]. Accordingly, the case where the segmentation cavity enlarges and the embryo developes into a hollow blastosphere may, under the circumstances, be simply described as the case where that outflow or exudation from the cells of the blastoderm is directed on the whole inwards.

The physical forces involved in the invagination of the cell-layer to form the gastrula have been repeatedly discussed[385], but the true explanation seems as yet to be by no means clear. The case, however, is probably not a very difficult one, provided that we may assume a difference of osmotic pressure at the two poles of the blastosphere, that is to say between the cells which are being differentiated into outer and inner, into epiblast and hypoblast. It is plain that a blastosphere, or hollow vesicle bounded by a layer of vesicles, is under very different physical conditions from a single, simple vesicle or bubble. The blastosphere has no effective surface tension of its own, such as to exert pressure on {345} its contents or bring the whole into a spherical form; nor will local variations of surface energy be directly capable of affecting the form of the system. But if the substance of our blastosphere be sufficiently viscous, then osmotic forces may set up currents which, reacting on the external fluid pressure, may easily cause modifications of shape; and the particular case of invagination itself will not be difficult to account for on this assumption of non-uniform exudation and imbibition.

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