On Growth and Form

Each evolution of form, from simpler to more complex, is deemed to have been attended by an advantage to the organism, an enhancement of its chances of survival or perpetuation; hence the historically older forms are, on the whole, structurally the simpler; or conversely the simpler forms, such as the simple sphere, were the first to come into being in primeval seas; and finally, the gradual development and increasing complication of the individual within its own lifetime is held to be at least a partial recapitulation of the unknown history of its race and dynasty*.

We encounter many, difficulties when we try to extend such concepts as these to the Foraminifera. We are led for instance to assert, as Rhumbler does, that the increasing complexity of the shell, and of the manner in which one chamber is fitted on another, makes for advantage; and the particular advantage on which Rhumbler rests his argument is strength. Increase of strength, die Festigkeitssteigerung, is according to him the guiding principle in foraminiferal evolution, and marks the historic stages of their development in geologic time. But in days gone by I used to see the beach of a little Connemara bay bestrewn with millions upon millions of foraminiferal shells, simple Lagenae, less simple Nodosariae, more complex Rotalae: all drifted by wave and gentle current from their sea-cradle to their sandy grave: all lying bleached and dead: one more delicate than another, but all (or vast multitudes of them) perfect and unbroken. And so I am not inclined to believe that niceties of form affect the case very much: nor in general that foraminiferal life involves a struggle for existence wherein breakage is a constant danger to be averted, and increased strength an advantage to be ensuredt.

In the course of the same argument Rhumbler remarks that Foraminifera are absent from the coarse sands and gravels{, as Williamson indeed had observed many years ago: so averting, or * A difficulty arises in the case of forms (like Peneroplis) where the young shell appears to be more complex than the old, the first formed portion being closely coiled while the later additions become straight and simple: “die biformen Arten verhalten sich, kurz gesagt. gerade umgekehrt als man nach dem biogenetischen Grundgesetz erwarten sollte,” Rhumbler, op. cit., p. 33 ete.

7 “Das Festigkeitsprinzip als Movens der Weiterentwicklung ist zu interessant und fiir die Autstellung meines Systems zu wichtig um die Frage unerértert zu lassen, warum diese Bevorziigung der Festigkeit stattgefunden hat. Meiner Ansicht nach lautet die Antwort auf diese Frage einfach, weil die Foraminiferen meistens unter Verhaltnissen leben, die ihre Schalen in hohem Grade der Gefahr des Zerbrechens aussetzen; es muss also eine fortwahrende Auslese des Festeren stattfinden,” Rhumbler, op. cit., p. 22.

t “Die Foraminiferen kiesige oder grobsandige Gebiete des Meeresbodens nicht lieben, u.s.w.”: where the last two words have no particular meaning, save only that (as M. Aurelius says) “of things that use to be, we say commonly that they love to be.” at least escaping, the dangers of concussion. But this is after all a very simple matter of mechanical analysis. The coarseness or fineness of the sediment on the sea-bottom is a measure of the current: where the current is strong the larger stones are washed clean, where there is perfect stillness the finest mud settles down; and the light, fragile shells of the Foraminifera find their appropriate place, like every other graded sediment, in this spontaneous order of lixiviation.

The theorem of Organic Evolution is one thing; the problem of deciphering the lines of evolution, the order of phylogeny, the degrees of relationship and consanguinity, is quite another. Among the higher organisms we arrive at conclusions regarding these things by weighing much circumstantial evidence, by dealing with the resultant of many variations, and by considering the probability or improbability of many coincidences of cause and effect; but even then our conclusions are at best uncertain, our judgments are continually open to revision and subject to appeal, and all the proof and confirmation we can ever have is that which comes from the direct, but fragmentary evidence of palaeontology*.

But in so far as forms can be shewn to depend on the play of physical forces, and the variations of form to be directly due to simple quantitative variations in these, just so far are we thrown back on our guard before the biological conception of consanguinity, and compelled to revise the vague canons which connect classification with phylogeny. The physicist explains in terms of the properties of matter, and classifies according to a mathematical analysis, all the drops and forms of drops and associations of drops, all the kinds of froth and foam, which he may discover among inanimate things; and his task ends there. But when such forms, such conformations and configurations, occur among lwing things, then at once the biologist introduces his concepts of heredity, of historical evolution, of succession in time, of recapitulation of remote ancestry in individual growth, of common origin (unless contradicted by direct evidence) of similar forms remotely separated by geographic space or geologic time, of fitness for a function, of

* Tn regard to the Foraminifera, “‘die Palaeontologie lasst uns leider an Anfang der Stammesgeschichte fast ginzlich im Stiche,” Rhumbler, op. cii., p. 14. adaptation to an environment, of higher and lower, of “better” and “worse.” This is the fundamental difference between the “explanations” of the physicist and those of the biologist. In the order of physical and mathematical complexity there is no question of the sequence of historic time. The forces that bring about the sphere, the cylinder or the ellipsoid are the same yesterday and to-morrow. A snow-crystal is the same to-day as when the first snows fell. The physical forces which mould the forms of Orbulina, of Astrorhiza, of Lagena or of Nodosaria to-day were still the same, and for aught we have reason to believe the physical conditions under which they worked were not appreciably different, in that yesterday which at call the Cretaceous epoch ; or, for aught we know, throughout all that duration of time which is marked, but not measured, by the geological record.

In a word, the minuteness of our organism brings its conformation as a whole within the range of the molecular forces; the laws of its growth and form appear to le on simple lines; what Bergson calls* the “ideal kinship” is plain and certain, but the “material affiliation” is problematic and obscure; and, in the end and upshot, it seems to me by no means certain that the biologist’s usual mode of reasoning is appropriate to the case, or that the concept of continuous historical evolution must necessarily, or may safely and legitimately, be employed.

* The evolutionist theory, as Bergson puts it, “‘consists above all in establishing relations of ideal kinship, and in maintaining that wherever there is this relation of, so to speak, logical affiliation between forms, there is also a relation of chronological succession between the species in which these forms are materialised” : Creative Evolution, 1911, p. 26. Cf. supra, p. 251. We have had so much to say on the subject of shell-spirals that we must deal briefly with the analogous problems which are presented by the horns of sheep, goats, antelopes and other horned quadrupeds; and all the more, because these horn-spirals are on the whole less symmetrical, less easy of measurement than those of the shell, and in other ways also are less easy of investigation. Let us dispense altogether in this case with mathematics ; and be content with a very simple account of the configuration of a horn.

There are three types of horn which deserve separate consideration: firstly, the horn of the rhinoceros; secondly the horns of the sheep, the goat, the ox or the antelope, that is to say, of the so-called hollow-horned ruminants; and thirdly, the solid bony horns, or “antlers,” which are characteristic of the deer. The horn of the rhinoceros presents no difficulty. It is physiologically equivalent to a mass of consolidated hairs, and, like ordinary hair, it consists of non-living or “formed” material, continually added to by the living tissues at its base. In section, that is to say in the form of its “generating curve,” the horn is approximately elliptical, with the long axis fore-and-aft, or, in some species, nearly circular. Its longitudinal growth proceeds with a maximum velocity anteriorly, and a minimum posteriorly ; and the ratio of these velocities being constant, the horn curves into the form of a logarithmic spiral in the manner that we have already studied. The spiral is of small angle, but in the longerhorned species, such as the great white rhinoceros (Ceratorhinus), the spiral form is distinctly to be recognised. As the horn

occupies a median position on the head,—a position, that is to say, of symmetry in respect to the field of force on either side,—there is no tendency towards a lateral twist, and the horn accordingly develops as a plane logarithmic spiral. When two horns coexist, the hinder one is much the smaller of the two: which is as much as to say that the force, or rate, of growth diminishes as we pass backwards, just as it does within the limits of the single horn. And accordingly, while both horns have essentially the same shape, the spiral curvature is less manifest in the second one, simply by reason of its comparative shortness.

The paired horns of the ordinary hollow-horned ruminants, such as the sheep or the goat, grow under conditions which are in some respects similar, but which differ in other and important respects from the conditions under which the horn grows in the rhinoceros. As regards its structure, the entire horn now consists of a bony core with a covering of skin; the inner, or dermal, layer of the latter is richly supplied with nutrient blood-vessels, while the outer layer, or epidermis, develops the fibrous or chitinous material, chemically and morphologically akin to a mass of cemented or consolidated hairs, which constitutes the “sheath” of the horn. A zone of active growth at the base of the horn keeps adding to this sheath, ring by ring, and the specific form of this annular zone is, accordingly, that of the “ generating curve” of the horn. Each horn no longer lies, as it does in the rhinoceros, in the plane of symmetry of the animal of which it forms a part; and the limited field of force concerned in the genesis and growth of the horn is bound, accordingly, to be more or less laterally asymmetrical. But the two horns are in symmetry one with another; they form “conjugate” spirals, one being the “ mirror-image” of the other. Just as in the hairy coat of the animal each hair, on either side of the median “parting,” tends to have a certain definite direction of its own axis, inclined away from the median axial plane of the whole system, so is it both with the bony core of the horn and with the consolidated mass of hairs or hair-like substance which constitutes its sheath; the primary axis of the horn is more or less inclined to, and may even be nearly perpendicular to, the axial plane of the animal.

The growth of the horny sheath is not continuous, but more or less definitely periodic: sometimes, as in the sheep, this periodicity is particularly well-marked, and causes the horny sheath to be composed of a series of all but separate rings, which are supposed to be formed year by year, and so to record the age of the animal*. Just as we sought for the true generating curve in the orifice, or “lip,” of the molluscan shell, so we might be apt to assume that in the spiral horn the generating curve corresponded to the lip or margin of one of the horny rings or annuli. This annular margin, or boundary of the ring, is usually a smuous curve, not lying in a plane, but such as would form the boundary of an anticlastic surface of great complexity: to the meaning and origin of which phenomenon we shall return presently. But, as we have already seen in the case of the molluscan shell, the complexities of the lip itself, or of the corresponding lines of growth upon the shell, need not concern us in our study of the development of the spiral: inasmuch as we may substitute for these actual boundary lines, their “trace,” or projection on a plane perpendicular to the axis—in other words the simple outline of a transverse section of the whorl. In the horn, this transverse section is often circular or nearly so, as in the oxen and many antelopes: it now and then becomes of somewhat complicated polygonal outline, as in a highland ram; but in many antelopes, and in most of the sheep, the outline is that of an isosceles, or sometimes nearly equilateral triangle, a form which is typically displayed, for instance, in Ovis Ammon. The horn in this latter case is a trihedral prism, whose three faces are, (1) an upper, or frontal face, in continuation of the plane of the frontal bone; (2) an outer, or orbital, starting from the upper margin of the orbit; and (3) an inner, or “nuchal,” abutting on the parietal boney. Along these three faces, and their corresponding angles or edges, we can trace in the fibrous substance of the horn a series of homologous spirals, such as we

* In the case of the ram’s horn, the assumption that the rings are annual is probably justified. In cattle they are much less conspicuous, but are sometimes well-marked in the cow; and in Sweden they are then called “calf-rings,” from a belief that they record the number of offspring. That is to say, the growth of the horn is supposed to be retarded during gestation, and to be accelerated after parturition, when superfluous nourishment seeks a new outlet. (Cf. Lénnberg, P.Z.S., p. 689, 1900.)

have called in a preceding chapter the “ensemble of generating spirals” which constitute the surface. In some few cases, of which the male musk ox is one of the most notable, the horn is not developed in a continuous spiral curve. It changes its shape as growth proceeds; and this, as we have seen, is enough to show that it does not constitute a logarithmic spiral. The reason is that the bony exostoses, or horn-cores, about which the horny sheath is shaped and moulded, neither grow continuously nor even remain of constant size after attaining their full growth. But as the horns grow heavy the bony core is bent downwards by their weight, and so guides the

Fig. 318. Diagram of Ram’s horns. (After Sir Vincent Brooke, from P.Z.8.) a, frontal; b, orbital; c, nuchal surface. growth of the horn in a new direction. Moreover as age advances, the horn-core is further weakened and to a great extent absorbed : and the horny sheath or horn proper, deprived of its support, continues to grow, but in a flattened curve very different from its original spiral*. The chamois is a somewhat analogous case. Here the terminal, or oldest, part of the horn is curved; it tends to assume a spiral form, though from its comparative shortness it seems merely to be bent into a hook. But later on, the bony core within, as it grows and strengthens, stiffens the horn, and guides it into a straighter course or form. The same phenomenon

of change of curvature, manifesting itself at the time when, or the place where, the horn is freed from the support of the internal core, 1s seen in a good many other antelopes (such as the hartebeest) and in many buffaloes; and the cases where it is most manifest appear to be those where the bony core is relatively short, or relatively weak. Fig. 319. Head of Arabian Wild Goat, Capra sinaitica. (After Sclater, from P.Z.S.) But in the great majority of horns, we have-no difficulty in recognising a continuous logarithmic spiral, nor in referring it, as before, to an unequal rate of growth (parallel to the axis) on two opposite sides of the horn, the inequality maintaining a constant ratio as long as growth proceeds. In certain antelopes, such as the gemsbok, the spiral angle is very small, or in other words the horn is very nearly straight; in other species of the same genus Oryx, such as the Beisa antelope and the Leucoryx, a gentle

curve (not unlike though generally less than that of a Dentalium shell) is evident; and the spiral angle, according to the few measurements I have made, is found to measure from about 20° to nearly 40°. In some of the large wild goats, such as the Scinde wild goat, we have a beautiful logarithmic spiral, with a constant angle of rather less than 70°; and we may easily arrange a series of forms, such for example as the Siberian ibex, the mouftion, Ovis Ammon, etc., and ending with the long-horned Highland ram: in which, as we pass from one to another, we recognise precisely homologous spirals, with an increasing angular constant, the spiral angle beg, for instance, about 75° or rather less in Ovis Ammon, and in the Highland ram a very little more. We have already seen that in the neighbourhood of 70° or 80° a small change of angle makes a marked difference in the appearance of the spire; and we know also that the actual length of the horn makes a very striking difference, for the spiral becomes especially conspicuous to the eye when a horn or shell is long enough to shew several whorls, or at least a considerable part of one entire whorl.

Even in the simplest cases, such as the wild goats, the spiral is never (strictly speaking) a plane or discoid spiral: but in greater or less degree there is always superposed upon the plane logarithmic spiral a helical spiral in space. Sometimes the latter is scarcely apparent, for the helical curvature is comparatively small, and the horn (though long, as in the said wild goats) is not nearly long enough to shew a complete convolution: at other times, as in the ram, and still better in many antelopes, such as the koodoo, the helicoid or corkscrew curve of the horn is its most characteristic feature.

Accordingly we may study, as in the molluscan shell, the helicoid component of the spire—in other words the variation in what we have called (on p. 555) the angle 6. This factor it is which, more than the constant angle of the logarithmic spiral, imparts a characteristic appearance to the various species of sheep, for instance to the various closely allied species of Asiatic wild sheep, or Argali. In all of these the constant angle of the logarithmic spiral is very much the same, but the shearing component differs greatly. And thus the long drawn out horns of

Ovis Poli, four feet or more from tip to tip, differ conspicuously from those of Ovis Ammon or O. hodgsoni, in which a very similar logarithmic spiral is wound (as it were) round a much blunter cone. The ram’s horn then, like the snail’s shell, is a curve of double curvature, in which one component has imposed upon the structure a plane logarithmic spiral, and the other has produced a continuous displacement, or “shear,” proportionate in magnitude to, and perpendicular or otherwise inclined in direction to, the axis of the former spiral curvature. The result is precisely analogous to that which we have studied in the snail and other spiral univalves ; but while the form, and therefore the resultant forces, are similar, the original distribution of force is not the same: for we have not here, as we had in the snail-shell, a “columellar” muscle, to introduce the component acting in the direction of the axis. We have, it is true, the central bony core, which in part performs an analogous function; but the main phenomenon here is apparently a complex distribution of rates of growth, perpendicular to the plane of the generating curve.

Let us continue to dispense with mathematics, for the mathematical treatment of a curve of double curvature is never very simple, and let us deal with the matter by experiment. We have seen that the generating curve, or transverse section, of a typical ram’s horn is triangular in form. Measuring (along the curve of the horn) the length of the three edges of the trihedral structure in a specimen of Ovis Ammon, and calling them respectively the outer, inner, and hinder edges (from their position at the base of the horn, relatively to the skull), I find the outer edge to measure 80cm., the inner 74cm., and the posterior 45cm.; let us say that, roughly, they are in the ratio of 9:8:5. Then, if we make a number of little cardboard triangles, equip each with three little legs (I make them of cork), whose relative lengths are as 9:8: 5, and pile them up and stick them all together, we straightway build up a curve of double curvature precisely analogous to the ram’s horn: except only that, in this first approximation, we have not allowed for the gradual increment (or decrement) of the triangular surfaces, that is to say, for the tapering of the horn due to the growth in its own plane of the generating curve.

In this case then, and in most other trihedral or three-sided horns, one of the three components, or three unequal velocities of growth, is of relatively small magnitude, but the other two are nearly equal one to the other. It would involve but little change for these latter to become precisely equal; and again but little to turn the balance of inequality the other way. But the immediate consequence of this altered ratio of growth would be that the horn would appear to wind the other way, as it does in the antelopes, and also in certain goats, e.g. the markhor, Capra falconeri.

For these two opposite directions of twist Dr Wherry has introduced a convenient nomenclature. When the horn winds so that we follow it frem base to apex in the direction of the hands of a watch, it is customary to call it a “left-handed” spiral. Such a spiral we have in the horn on the left-hand side ofaram’shead. Accordingly, Dr Wherry calls the condition homonymous, where, as in the sheep, a right-handed spiral is on the right side of the head, and a left-handed spiral on the left side; while he calls the opposite condition heteronymous, as we have it in the antelopes, where the right-handed twist is on the left side of the head, and the left-handed twist on the right-hand side. Among the goats, we may have either condition. Thus the domestic and most of the wild goats agree with the sheep; but in the markhor the twisted horns are heteronymous, as in the antelopes. The difference, as we have seen, is easily explained; and (very much as in the case of our opposite spirals in the apple-snail, referred to on p. 560), it has no very deep importance.

Summarised then, in a very few words, the argument by which we account for the spiral conformation of the horn is as follows: The horn elongates by dint of continual growth within a narrow zone, or annulus, at its base. If the rate of growth be identical on all sides of this zone, the horn will grow straight; if it be greater on one side than on the other, the horn will become curved : and it probably will be greater on one side than on the other, because each single horn occupies an unsymmetrical field with reference to the plane of symmetry of the animal. If the maximal and minimal velocities of growth be precisely at opposite sides of the zone of growth, the resultant spiral will be a plane spiral: but if they be not precisely or diametrically opposite, then the spiral will be a spiral in space, with a winding or helical component; and it is by no means likely that the maximum and minimum will occur at precisely opposite ends of a diameter, for

no such plane of symmetry is manifested in the field of force to which the growing annulus corresponds or appertains. Now we must carefully remember that the rates of growth of which we are here speaking are the net rates of longitudinal increment, in which increment the activity of the living cells in the zone of growth at the base of the horn is only one (though it is the fundamental) factor. In other words, if the horny sheath were continually being added to with equal rapidity all round its zone of active growth, but at the same time had its elongation more retarded on one side than the other (prior to its complete solidification) by varying degrees of adhesion or membranous attachment to the bone core within, then the net result would be a spiral curve precisely such as would have arisen from initial inequalities in the rate of growth itself. It seems highly probable that this is a very important factor, and sometimes even the chief factor in the case. The same phenomenon of attachment to the bony core, and the consequent friction or retardation with which the sheath slides over its surface, will lead to various subsidiary phenomena: among others to the presence of transverse folds or corrugations upon the horn, and to their unequal distribution upon its several faces or edges. And while it is perfectly true that nearly all the characters of the horn can, be accounted for by unequal velocities of longitudinal growth upon its different sides, it is also plain that the actual field of force is a very complicated one indeed. For example, we can easily see that (at least in the great majority of cases) the direction of growth of the horny fibres of the sheath is by no means parallel to the axis of the core within; accordingly these fibres will tend to wind in a system of helicoid curves around the core, and not only this helicuid twist but any other tendency to spira] curvature on the part of the sheath will tend to be opposed or modified by the resistance of the core within.

But on the other hand living bone is a very plastic structure, and yields easily though slowly to any forces tending to its deformation; and so, to a considerable extent, the bony core itself will tend to be modelled by the curvature which the growing sheath assumes, and the final result will be determined by an equilibrium between these two systems of forces. While it is not very safe, perhaps, to lay down any general tule as to what horns are more, and what are less spirally curved, I think it may be said that, on the whole, the thicker the horn, the greater is its spiral curvature. It is the slender horns, of such forms as the Beisa antelope, which are gently curved, and it is the robust horns of goats or of sheep in which the curvature is more pronounced. Other things being the same, this is what we should expect to find; for it is where the transverse section of the horn is large that we may expect to find the more marked differences in the intensity of the field of force, whether of active growth or of retardation, on opposite sides or in different sectors thereof.

But there is yet another and a very remarkable phenomenon which we may discern in the growth of a horn, when it takes the form of a curve of double curvature, namely, an effect of torsional strain; and this it is which gives rise to the sinuous “lines of growth,” or sinuous boundaries of the separate horny rings, of which we have already spoken. It is not at first sight obvious that a mechanical strain of torsion is necessarily involved in the growth of the horn. In our experimental illustration (p. 618), we built up a twisted coil of separate elements, and no torsional strain attended the development of the system. So would it be if the horny sheath grew by successive annular increments, free save for their relation to one another, and having no attachment to the solid core within. But as a matter of fact there is

such an attachment, by subcutaneous connective tissue, to the bony core; and accordingly a torsional strain will be set up in the growing horny sheath, again provided that the forces of growth therein be directed more or less obliquely to the axis of the core; for a “couple” is thus introduced, giving rise to a strain which the sheath would not experience were it free (so to speak) to slip along, impelled only by the pressure of its own growth from below. And furthermore, the successive small increments of the growing horn (that is to say, of the horny sheath) are not instantaneously converted from living to solid and rigid substance; but there is an intermediate stage, probably long-continued, during which the new-formed horny substance in the neighbourhood of the zone of active growth is still plastic and capable of deformation. Now we know, from the celebrated experiments of St Venant*, that in the torsion of an elastic body, other than a cylinder of circular section, a very remarkable state of strain is introduced. If the body be thus cylindrical (whether solid or hollow), then a twist leaves each circular section unchanged, in dimensions and in figure. But in all other cases, such as an elliptic rod or a prism of any particular sectional form, forces are introduced which act parallel to the axis of the structure, and which warp each section into a complex anticlastic surface. Thus in the case of a triangular and equilateral prism, such as is shewn in section in Fig. 321, if the part of the rod represented in the section be twisted by a force acting in the direction of the arrow, then the originally plane section will be warped as indicated in the diagram :—where the full contour-lines represent elevation above, and the dotted lines represent depression below, the original level.